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Full text of "Bulletin of the British Museum (Natural History) Entomology"

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Bulletin of the 

British Museum (Natural History) 



Entomology series Vol 56 1 987-88 



t»U8. f . 



9 At 



British Museum (Natural History) 
London 1988 



Dates of publication of the parts 

Nol 30 July 1987 

No 2 29 October 1987 

No 3 17 December 1987 

No 4 . . ..• 28 April 1988 



ISSN 0524-6431 



Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 



Contents 

Entomology Volume 56 

Page 
No 1 The legume-feeding psyllids (Homoptera) of the west Palaearctic 

Region. Byl.D. Hodkinson&D. Hollis 1 

No 2 A review of the Malvales-feeding psyllid family Carsidaridae 

(Homoptera). By D. Hollis 87 

No 3 A review of the Rhadalinae ( = Aplocneminae) (Coleoptera: 

Melyridae). By Enid R. Peacock 129 

No 4 A revision of some Afrotropical genera of Eucoilidae (Hymenoptera). 

By J. Quinlan 171 



S.\. 



Bulletin of the 

British Museum (Natural History) 



IDA' 

3/ 

pf 



The legume-feeding psyllids (Homoptera) 
of the west Palaearctic Region 

I. D. Hodkinson & D. Hollis 



Entomology series 
Vol 56 No 1 



30 July 1987 



The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, 
and an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
ever-growing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
available separately, and individually priced. Volumes contain about 300 pages and several 
volumes may appear within a calendar year. Subscriptions may be placed for one or more of 
the series on either an Annual or Per Volume basis. Prices vary according to the contents of 
the individual parts. Orders and enquiries should be sent to: 



Publications Sales, 

British Museum (Natural History), 
Cromwell Road, 

London SW7 5BD, 
England. 



World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) 



© British Museum (Natural History), 1987 



The Entomology series is produced under the editorship of the 

Keeper of Entomology: Laurence A. Mound 

Entomology Publications Manager: W. Gerald Tremewan 



ISBN 565 06028 7 
ISSN 0524-6431 



British Museum (Natural History) 
Cromwell Road 
London SW7 5BD 




Entomology series 
Vol 56 No lpp 1-86 



Issued 30 July 1987 



The legume-feeding psyllids (Homoptera) of the west 
Palaearctic Region 

I. D. Hodkinson 

Department of Biology, Liverpool Polytechnic, Byrom Street, Liverpool L3 3AF 

D. Hollis 

Department of Entomology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Contents 

Synopsis 1 

Introduction 2 

Material, methods and terminology 3 

Acknowledgements 3 

Key to west Palaearctic genera of Arytaininae and Acizziinae 3 

Acizziinae White & Hodkinson 4 

Acizzia Heslop-Harrison 4 . 

Key to species 4 

Arytaininae Crawford 7 

Cyamophila Loginova 7 

Key to species 8 

Arytaina Forster 10 

Key to species 11 

Livilla Curtis 19 

Key to species 20 

Arytainilla Loginova 41 

Discussion 43 

References 44 

Index to host plants 84 

Index 84 

Synopsis 

Plant species of the papilionoid legume tribe, the Genisteae, have radiated considerably in the Mediter- 
ranean Basin and are exploited, as hosts, by psyllids of the subfamily Arytaininae; other leguminous hosts 
exploited by this group of psyllids belong to the papilionoid tribes Galegeae, Trifoleae, Loteae and 
Sophoreae; species of the related psyllid subfamily Acizziinae develop on host species in the mimosoid 
genera Acacia and Albizia. 

The taxonomy of the psyllid groups Arytaininae and Acizziinae in the west Palaearctic Region is revised; 
the Arytaininae is considered to be a paraphyletic group but is retained for convenience; the Acizzinae is 
considered to be a monophyletic group. Emphasis is placed on characters of the genitalia in delimiting 
genera and five are recognised: Arytaina s. str. (11 species), Livilla (31 species), Cyamophila (4 species) 
and Arytainilla s.l. (21 species) in the Arytaininae, and Acizzia (4 species) in the Acizzinae. The genera 
Floria syn. n. and Alloeoneura syn. n. are synonymised with Livilla; Amblyrhina syn. n. is synonymised 
with Arytaina. Diagnostic keys are provided for the genera and identification keys are given for their 
included species except those of Arytainilla, a problematical genus that is discussed only in general terms. 
Sixteen new species are described; 3 new species-synonymies, 48 new combinations and one new name are 
proposed; one neotype and 5 lecto types are designated; Peripsyllopsis is reinstated as a good genus. 

The pattern of host/insect relationships between these psyllids and their leguminous hosts is examined 
and it is concluded that, with the present level of data, an understanding of psyllid interrelationships does 
not clarify the problems associated with the classification and evolution of the Genisteae. 



Bull. Br. Mus. nat. Hist. (Ent.) 56 (1): 1-86 Issued 30 July 1987 



D. HODKINSON & D. HOLLIS 



Introduction 

The Mediterranean Basin and the surrounding land areas of the west Palaearctic Region, 
together with Macaronesia, form an extensive area of endemism and diversity for a tribe of 
papilionoid leguminous plants, the Genisteae. This tribe has been successfully exploited as hosts 
by over 60 known species of jumping plant-lice, or psyllids, previously referred to the subfamily 
Arytaininae (Hodkinson, 1980). Other psyllids in this group colonise legumes in the papilionoid 
tribes Galegeae, Trifoleae, Loteae and Sophoreae and in the mimosoid genera Acacia and 
Albizia. 

The detailed biologies of most of these psyllids are unknown. The free-living nymphs usually 
feed on the actively growing shoots of their hosts and can often reach high densities. Watmough 
(1968a, b) recorded peak densities of 6466 and 166 nymphs per 100 g of shoot material for 
Arytainilla spartiophila and Arytaina genistae respectively, on Cytisus scoparius in Britain. 
Acizzia uncatoides has caused severe damage to acacia trees where it has been introduced into 
California, France and Hawaii (Koehler et al., 1966; Bain et al, 1976; Leeper & Beardsley, 
1976). Life history strategies and generation times appear to vary between species. Arytainilla 
spartiophila, in Britain, has a single generation per year with six months spent as an overwinter- 
ing diapause egg. By contrast, Arytaina genistae overwinters as an adult and undergoes two or 
three generations per year on the same host. 

The Arytaininae were reviewed by Loginova (1976a; 1977), who divided the group into two 
tribes. The genera Arytaina, Amblyrhina, Alloeoneura, Floria, Livilla and the central Asian 
Astragalita were included in the Arytainini; Cyamophila, Acizzia, Arytainilla and the North 
American Amorphicola were included in the Cyamophilini. The monotypic genus Pseudacan- 
thopsylla was omitted. White & Hodkinson (1985) placed Acizzia in a separate subfamily, the 
Acizziinae, primarily on the basis of the form of the male proctiger, and expressed uncertainty as 
to the true position of Cyamophila. 

Examination of species within the above genera, together with new material, leads us to 
conclude that the major reorganisation of species within genera, proposed in this paper, is 
necessary if the generic classification is to reflect natural groupings. Previous classifications have 
been based largely on primitive characters and the resulting taxa have been heterogeneous and 
ill-defined, and closely related species have often been placed in separate genera. We have 
attempted here to recognise what we believe are monophyletic genera or groups based mainly 
on characters of the male genitalia. 

Six legume-feeding genera are recognised in the west Palaearctic Region: Arytaina s. str., 
Livilla, Cyamophila, Arytainilla, Acizzia and Pseudacanthopsylla. Amblyrhina, Floria and 
Alloeoneura are placed in synonymy. Arytainilla is retained as an heterogeneous genus but we 
recognise that it may be polyphyletic. Pseudacanthopsylla shows affinities with undescribed 
Afrotropical groups and is excluded from further consideration. We recognise two separate 
subfamilies, the Acizzinae which contains Acizzia, and the Arytaininae which contains Ary- 
taina, Livilla, Cyamophila and Arytainilla. The Acizziinae are defined by characters of the male 
genitalia. We are unable, however, to find synapomorphies that would define the Arytaininae 
reliably and separate it from the closest subfamilies, the Psyllinae and Ciriacreminae. Loginova 
(1976a) gave a lengthy description of the group, including both adult and larval morphology. A 
detailed study of these characters has shown that none is present in all Arytainines and absent 
from other groups within the Psyllidae, and several are either primitive or subject to homoplasy. 
Furthermore no previously unused characters could be found. We therefore regard the 
Arytaininae as paraphyletic with respect to the Ciriacreminae and Psyllinae but, nevertheless, 
we retain the group as a convenient and widely understood entity embracing groups of psyllids 
that feed on legumes. 

This paper contains a complete account of Arytaina and Livilla, and accounts of those species 
of Acizzia and Cyamophila that occur in the west Palaearctic Region. Arytainilla is dealt with in 
less detail as the species have been well-described and figured by Sulc (1907; 1910a, b) and 
Loginova (1972a; 19766). The pattern of host/psyllid relationships between the groups of 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 3 

Genisteae genera and psyllid genera is also examined to see if there is any degree of congruence 
between their respective classifications. 

Materials, methods and terminology 

A detailed explanation of the terminology used in this paper is given by Hodkinson & White (1979/?) and 
White & Hodkinson (1985). The following abbreviations are used to indicate the institutions in which 
material is deposited: British Museum (Natural History), London (BMNH); Museum National d'Histoire 
Naturelle, Paris (MNHN); Musee de Zoologie in Lausanne (MZL); Zoological Institute, Academy of 
Sciences of the U.S.S.R., Leningrad (ZI); Moravian Museum, Brno (MM); Naturhistorisches Museum, 
Vienna (NM); Naturhistoriches Museum, Basel (NMB); Zoological Museum, University of Helsinki 
(ZMU); Termeszettudomanyi Muzeum, Budapest (TM), Zoological Institute, Polish Academy of 
Sciences, Warsaw (IZPAN); Entomological Institute Eidgenossische Technische Hochschule, Zurich 
(ETH); Museo Civico di Storia Naturale, Genova (MCSN); New Zealand Arthropod Collection, 
Department of Scientific and Industrial Research, Auckland (NZAC); University Museum, Melbourne 
(UMM); I. D. Hodkinson collection (IDH); P. Lauterer collection (PL); D. Burckhardt collection (DB). 
All drawings and measurements are made from slide-mounted material. 

Acknowledgements 

We thank the authorities and curators at the institutions listed above for the loan of material . We also thank 
Dr P. Lauterer, Dr D. Burckhardt, Dr S. M. Klimaszewski, Dr C. Conci and the late Dr M. M. Loginova 
for the loan of interesting and often undescribed material. 

Key to west Palaearctic genera of Ary taininae and Acizziinae 

1 Male proctiger (Figs 9, 15) with a well-developed posterior lobe. On Acacia or Albizia. 

(Genal cones shorter than the vertex, triangular or broadly rounded, rarely enveloping the 
frons. Antennae approximately 1-3-1-8 times head width. Propleurites broad, divided by a 
vertical suture. Forewing oblong-oval, with or without pattern but with a distinct costal break 
and pterostigma; cell cw la high. Basal metatarsus with 1 or 2 thick black spurs. Male paramere 
of various shapes) . (Subfamily ACIZZIINAE) ACIZZIA Heslop-Harrison (p. 4) 

- Male proctiger (Figs 23, 68, 167) simple, without a well-developed posterior lobe (Subfamily 

ARYTAININAE) 2 

2 Male paramere (Figs 23, 39, 75, 82) broad, more or less parallel-sided with a truncate apex which 

is extended anteriorly in the form of a pronounced tooth 3 

- Male paramere not of this form, either slender and gradually narrowed to apex which bears a 

slender inner anteriorly curving tooth (Figs 167, 176, 189, 195), or of more irregular shape 4 

3 Forewing (Figs 21, 26) usually oblong-oval, with both a well-developed pterostigma and a costal 

break; cell cw la high. Propleurites (Fig. 261) narrow, divided by a diagonal suture, pronotum 
deflexed downwards. Basal metatarsus with two thick black spurs. Not on Genisteae. 
(Genal cones shorter than the vertex, broadly rounded, deflexed from the plane of the vertex. 
Eyes less convex than in other Arytrainine genera. Antennae relatively stout, approximately 
1-2-1-6 times the head width. Forewing with or without pattern). 

CYAMOPHILA Loginova (p. 7) 

- Forewing (Figs 45, 49, 57) usually more oval, or more narrowly rounded at the apex, without 

well-developed pterostigma or costal break: cell cw la usually lower. Propleurites (Fig. 260) 
quadrate, divided by a vertical suture, pronotum flat. Basal metatarsus usually with one thick 
black spur. On host plants in the tribe Genisteae. 

(Genal cones shorter than the vertex, triangular, broadly rounded or truncate. Eyes promi- 
nent. Antennae variable, stout to slender, 1-1-2-3 times head width.) 

ARYTAINA F6rster(p. 10) 

4 Male paramere (Figs 167, 176, 189, 195) a simple narrow lamellar structure gradually tapering to 

the apex, which bears a small inner denticle, the apex of which is directed anteriorly. Forewing 
(Figs 103, 107, 109, 115, 125, 135) of variable shape, usually with a distinct colour pattern or 
coriaceous brown or amber-coloured (L. genistae is the exception). Pterostigma, if present, 
very short; costal break present or absent. Genal cones (Figs 104, 114, 124, 154) usually long 
and broad, in lateral view (Fig. 259) weakly separated from the vertex (exceptions are L. 
bivittata, L. cognata, L. burckhardti and L. vicina) (Figs 105, 109, 110, 166). Vertex and 



4 I. D. HODKINSON & D. HOLLIS 

thorax usually covered in long dense setae (much reduced in L. ulicis, L. vicina and L. 

bivittata). 

(Antennae slender, about 1-6-3-0 times width of the head. Eyes prominent. Pronotum flat; 

propleurites broad, divided by a vertical suture. Cell cw la of forewing usually low. Basal 

metatarsus with to 2 thick black spurs (usually 1)) LIVILLA Curtis (p. 19) 

- Male paramere not as above, often of more irregular shape or if evenly narrowed then much 

longer and thinner than those illustrated for Livilla. Forewing membrane without distinct 
pattern, occasionally with a pigment pattern confined to the veins, membrane usually clear, 
occasionally yellowish/orange but never coriaceous. Long pterostigma and costal break 
usually present (A. nubivaga and A. devia are the exceptions). Genal cones shorter, in lateral 
view usually separated from the vertex by a distinct step. Vertex and thorax not covered in 
long setae. 

(Antennae slender, elongate, approximately 1-4-2-4 times head width. Propleurites broad, 
divided by a vertical suture. Forewing usually oblong oval, cell cw la low. Basal metatarsus with 
one thick black spur.) AR YTAINILLA Loginova (p. 41) 

ACIZZIINAE White & Hodkinson 

Acizziinae White & Hodkinson, 1985: 271. 

The Acizziinae are separated from the Arytaininae/Psyllinae by the possession of a posteriorly lobed 
proctiger in the male. A single genus, Acizzia, occurs in the Palaearctic Region. 

ACIZZIA Heslop-Harrison 

Neopsylla Heslop-Harrison, 1949: 161; 1951: 417. Type-species: Psylla acaciae Maskell, 1894, by original 

designation. [Homonym of Ateopsy lla Wagner, 1903.] 
Acizzia Heslop-Harrison, 1961a: 417; Loginova, 1976a: 596; 1977: 577; White & Hodkinson, 1985: 271. 

[Replacement name for Neopsylla Heslop-Harrison.] 

Diagnosis. Male proctiger with a conspicuous posterior lobe which may or may not bear a secondary 
finger-like appendage (Figs 9, 15); forewing (Figs 1, 5) with a tapered pterostigma and a conspicuous costal 
break; cell cw la high and characteristically shaped, leaning towards the base of the wing; genal cones little 
deflexed, short, rounded or pyramidal, usually not completely enveloping the frons which bears the 
median ocellus (Figs 2, 8); basal metatarsus with 1 or 2 black spurs; apical segment of aedeagus often 
complex (Fig. 17); propleurites quadrate, divided by a vertical suture. Species feed on mimosoid legumes 
of the tribes Acacieae and Ingae, particularly Acacia and Albizia. 

Comments. Acizzia is a widely distributed genus of more than 30 described species found in Australia, New 
Zealand, the Old World tropics and extending through North Africa and the Middle East to the 
Mediterranean Basin. Four species occur in the area considered here, two of which are introduced from 
Australia. 

Key to species 

1 Forewing (Figs 5, 7) with pattern consisting of small diffuse spots primarily in the apical half. 

Male proctiger (Figs 15, 18) with long tubular apical portion and basal triangular lobe which 
bears a subsidiary finger-like projection (acaciaebaileyanae-group) 2 

- Forewing without pattern, or pattern consisting of brown clouds adjacent to the wing apex, 

primarily around veins R s , M 1+2 , M 3+4 and Cw la (Figs 2, 4). Male proctiger (Figs 9, 12) with 
single large triangular lobe without subsidiary pro j ections (hollisi-growp) 3 

2 Male proctiger (Fig. 18) with the tubular apical portion short and broad. Paramere as in Fig. 19. 

Apex of aedeagus (Fig. 20) bulbous. Female proctiger (Fig. 294) with dorsal margin evenly 
bluntly acute. Forewing pattern (Fig. 7) overlying areas of conspicuously thickened spinules. 
Smaller species, head width less than 0-47 mm acaciaebailevanae (Froggatt) (p. 5) 

- Male proctiger (Fig. 15) with tubular apical portion longer and narrower. Paramere as in Fig. 

16. Apex of aedeagus (Fig. 17) harpoon-shaped. Female proctiger (Fig. 293) with dorsal 
margin sinuous, apex slender. Forewing pattern (Fig. 5) not corresponding with areas of 
conspicuously thickened spinules. Larger species, head width more than 0-50 mm 

uncatoides (Ferris & Klyver) (p. 5) 

3 Forewing (Fig. 3) with apical brown clouds, apex broadly rounded. Male proctiger (Fig. 12) with 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 5 

a broadly round posterior lobe. Aedeagus (Fig. 14) with a straight bulbous apex. (Israel, on 

Acacia raddiana.) hollisi Burckhardt (p. 6) 

- Forewing (Fig. 1) without obvious colour pattern, apex narrowly rounded. Male proctiger (Fig. 
9) with more triangular posterior lobe. Aedeagus (Fig. 11) with reniform apex. (Egypt, on 
unknown host.) bicohrata (Samy) (p. 7) 

The acaciaebaileyanae-group 

Defined by characters in couplet 1 of key. This group comprises the introduced Australian species. 

Acizzia acaciaebaileyanae (Froggatt) 
(Figs 7, 8, 18-20,294) 

Psylla acaciae-baileyanae Froggatt, 1901: 257; Tuthill, 1952: 257. Lectotype d\ Australia, designated by 

Yen, 1977: 15 (ANIC). 
Arytaina acaciae-baileyanae (Froggatt) Pettey, 1924: 21. 
Psylla uncata Ferris & Klyver, 1932: 53. Syntypes d\ $, New Zealand (NZAC) [examined]. [Synony- 

mised by Tuthill, 1952: 91.] 
Neopsylla uncftjata (Ferris & Klyver) Heslop-Harrison, 1949: 162. [Mis-spelling.] 
Acizzia acaciaebaileyanae (Froggatt) Capener, 1970: 197; Loginova, 1977: 577; Hodkinson, 1983: 343; 

Morgan, 1984: 36. 
Psylla acaciaebaileyanae Froggatt; Yen, 1977: 15. 

Description. Coloration. General coloration of head and thorax dull orange-yellow with darker yellowish 
brown markings; abdominal sclerites yellowish brown, intersegmental membranes dull pale yellow; 
genitalia pale brown: forewing transparent with small irregular pale brown maculations; veins pale 
yellowish brown; antennae dull whitish yellow with apices of segments 3-7 infuscate and segments 8-10 
entirely dark brown; legs pale whitish yellow, often with faint dusky markings on the femora. 

Structure. Head (Fig. 8) robust, genal cones very short with broadly rounded apices. Forewing (Fig. 7) 
oblong oval, with a well-developed pterostigma and costal break; spinules present throughout all the cells, 
much larger in regions where the brown maculate pattern occurs than elsewhere; cell mi elongate; cell cw la 
high and strongly arched towards the wing base. Metatibia with 4 or 5 thick black apical spurs; basal 
metatarsus with 1 black spur. Male proctiger (Fig. 18) with a short and broad tubular apical projection and 
with a rounded posterior lobe bearing a subsidiary finger-like projection; paramere (Fig. 19) simple, broad 
basally, gradually tapering to rounded apex, with a small posteriorly directed subapical denticle on inner 
posterior margin; aedeagus (Fig. 20) with a bulbous apex and a long ductus ejaculatorius. Female 
terminalia (Fig. 294) with proctiger convex throughout dorsal margin, apex bluntly acute; ventral valve 
wedge shaped; circumanal pore ring simple, about one-third length of proctiger; ovipositor short and 
broad. 

Measurements. See Table 1. 

Material examined 

12 Cf , 13 $, Italy: on Acacia sp. (BMNH). 

Host plant. Acacia baileyana F.v.M. and A. podalyriaefoliae Cunn. 

Published distribution. Australia (Froggatt, 1901; Yen, 1977; Morgan, 1984); introduced to New 
Zealand (Tuthill, 1952; Ferris & Klyver, 1932), South Africa (Pettey, 1924; Capener, 1970), Italy 
(Rapisarda, 1985). 

Comments. This species is native to Australia and has recently been introduced into Italy on ornamental 
Acacia trees. It is distinguished from its close relative, A. uncatoides, by the shape of the genal cones and 
details of the male genitalia. 

Acizzia uncatoides (Ferris & Klyver) 
(Figs 5, 6, 15-17,293) 

Psylla uncatoides Ferris & Klyver, 1932: 53; Heslop-Harrison, 1949: 162; Tuthill, 1952: 89. Syntypes cf , $ , 

New Zealand (NZAC, BMNH) [examined]. 
Acizzia uncatoides (Ferris & Klyver) Loginova, 1977: 577; Hodkinson & White, 1981: 492; Hodkinson, 

1983: 343. 



6 I. D. HODKINSON & D. HOLLIS 

Description. Coloration. General body colour orange throughout with paler markings on dorsum of 
thorax; forewing membrane pale yellow to pale amber with orange-brown maculations, veins concolorous 
with membrane but appearing slightly darker; antennal segments 1-3 orange-yellow, the remainder 
brown; legs slightly paler yellow-brown, often with a slight infuscation of the femora. 

Structure. Head (Fig. 6) with vertex relatively long, genal cones triangular with rounded apices. 
Forewing (Fig. 5) oblong-oval, with well-developed pterostigma and costal break; spinules present 
throughout wing; cell m x elongate; cell cw la high and strongly arched towards the wing base. Metatibia with 
4 or 5 thick black spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 15) with long tubular apical 
portion and a rounded posterior lobe bearing a subsidiary finger-like projection; paramere (Fig. 16) broad, 
with apex deflexed posteriorly; aedeagus (Fig. 17) with harpoon-shaped apex. Female terminalia (Fig. 293) 
short and truncate with a relatively large circumanal pore ring. 

Measurements. See Table 1. 

Material examined 
Several hundred C\ $, Portugal, Israel, Chile (IDH, DB, USNM). 

Host plant. In Europe this species occurs on Acacia floribunda, while in Israel it is found on Acacia 
saligna. Elsewhere it has been recorded from Albizia lophantha, Acacia verniciflua, A. koaia, A. koa and 
A. confusa. 

Published distribution. Australia (Hodkinson, 1983); introduced to France (Bain et ai, 1976), Italy 
(Arzone & Vidano, 1985), U.S.A.: California (Jensen, 1957; Koehler etal , 1966; Madubunyi & Koehler, 
1974; Munro, 1965), Hawaii (Gagne, 1971; Leeper & Beardsley, 1973; 1976), New Zealand (Tuthill, 
1952), Mexico (Hodkinson & White, 1983). 

Comments. This species is native to Australia and has been introduced into several areas of the world 
where it has become a minor pest of ornamental or native acacias. Dr A. Yen (Australia) has compared our 
European material with material from Australia. A. uncatoides is separated from its close relative A. 
acaciaebaileyanae by the shape of the genal cones, the male proctiger, paramere and aedeagus. 



The hollisi-group 

Defined by characters in couplet 1 of key. This group includes the native North African or Middle East 
species. 

Acizzia hollisi Burckhardt 

(Figs 3, 4, 12-14,262) 

Acizzia hollisi Burckhardt, 1981: 216. Holotype d\ Saudi Arabia (NMB) [not examined]. 

Description. Coloration. General body coloration dark reddish brown throughout with paler markings on 
dorsum of thorax; intersegmental membranes of abdomen lighter orange-yellow; male proctiger often 
bright red, parameres whitish yellow; female terminalia dark brown; forewing membrane clear with brown 
colour pattern, veins brown; antennae yellowish brown, darkening towards apex; femora dark brown, 
remainder of legs pale yellowish brown. Immature specimens light orange-yellow. 

Structure. Head (Fig. 4) with vertex relatively short; genal cones shorter than vertex, triangular. 
Forewing (Fig. 3) oblong-oval, with well-developed pterostigma and costal break; spinules largely 
confined to apical half of wing and around vein Cu 2 ; cell cw la moderately strongly leaning towards wing 
base. Metatibia with 4 or 5 thick black spurs, basal metatarsus with 2 black spurs. Male proctiger (Fig. 12) 
with a broadly triangular posterior projection. Paramere (Fig. 13) with distinct notch in anterior margin; 
aedeagus (Fig. 14) with bulbous apex. Female terminalia (Fig. 262) relatively long and slender, with a 
relatively short anal pore ring. 

Measurements. See Table 1. 

Material examined 
3 cT , 3 $ (paratypes), 2 cf , 1 ?, Israel (BMNH, IDH). 

Host plant. Acacia raddiana. 

Published distribution. Saudi Arabia (Burckhardt, 1981; 1986), Israel (Burckhardt, 1981; Halperin et 
a/., 1982). 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 7 

Acizzia bicolorata (Samy) comb. n. 

(Figs 1,2, 9-11) 

Psylla bicolorata Samy, 1973: 451. Holotype cf , Egypt (Agr. Coll. Cairo) [not examined]. 

Description. Coloration. General body coloration orange-yellow throughout; forewing membrane pale 
yellow, veins concolorous; antennae orange-yellow basally, becoming brown apically; legs concolorous 
with body (based on a single specimen). According to Samy the general colour is bright brown with whitish 
markings on head and dorsum of the thorax. 

Structure. Head (Fig. 3) robust, genal cones shorter than vertex, with broadly rounded apices. Forewing 
(Fig. 2) with somewhat narrowly rounded apex; costal break and pterostigma well-developed; spinules 
confined to radular areas; cell cu ld high but less strongly displaced towards wing base than in uncatoides. 
Metatibia with 4 or 5 thick black spurs, basal metatarsus with 2 black spurs. Male proctiger (Fig. 10) with 
narrowly triangular posterior lobe; paramere (Fig. 11) simple, almost parallel-sided, apex pointing 
anteriorly; apex of aedeagus (Fig. 12) somewhat reniform. 

Measurements. See Table 1. 

Material examined 

1 cf (same data as holotype), Egypt: Oweinat, xi. 1937 (Kasim) (BMNH). 

Host plant. Unknown. 

Published distribution. Egypt (Samy, 1973). 

Comments . A . bicolorata is separated from the other west Palaearctic members of the genus by the lack of a 
forewing pattern. 

ARYTAININAE Crawford 

Arytainini Crawford, 1914: 106, in part; Heslop-Harrison, 1951: 417, in part. 

Arytaininae Crawford; Vondracek, 1957: 176; Loginova, 1976a: 589; 1977: 577; White & Hodkinson, 1985: 

272. 

The Arytaininae, as here constituted, is probably paraphyletic and overlaps with the subfamily Psyllinae. 
The group has been separated previously from the Psyllinae by the more quadrate propleurites, the 
presence of one, rather than two metatarsal spurs, the absent or rudimentary pterostigma and the genal 
cones which are usually in the same plane as the vertex, rather than being deflexed downwards. In addition , 
Arytainine species usually occur on Leguminosae whereas Psyllinae species occur usually on Rosaceae and 
a variety of other dicotyledonous families. However, Cyamophila, which appears to have a similar form of 
advanced paramere to Arytaina, has narrower propleurites, a fully developed pterostigma and the 2 
metatarsal spurs more typical of the Psyllinae. 

Together the Arytaininae and Psyllinae are defined by the rounded forewing with the venation pattern in 
which veins Cu x and M possess a common stem and vein Cu 2 terminates adjacent to vein Cu lb . The basal 
metatarsus usually possesses at least one spur, the genae are usually developed into cone-like processes and 
the 3rd antennal segment is the longest. The hind coxa also bears a well-developed meracanthus, and the 
male proctiger is simple and unipartite. 

CYAMOPHILA Loginova 

Cyamophila Loginova, 1976a: 596; 1977: 582; White & Hodkinson, 1985: 272. Type-species: Psylla fabra 
Loginova, 1964, by original designation. 

Diagnosis. Forewing (Figs 21, 26) with a well-developed pterostigma and costal break, cell cw la relatively 
high and arched towards base of wing; male paramere (Figs 24, 29) with apex truncate, formed of a large 
anteriorly directed denticle; basal metatarsus with 2 spurs; propleurites elongate, divided by a diagonal 
suture; genal cones (Figs 22, 27) short, in same plane as vertex, enveloping the median ocellus; eyes (Figs 
22, 27) less bulbous than in many other genera; antennae relatively stout, usually less than 1-5 x head 
width. 

Comments. Cyamophila is primarily a Palaearctic genus comprised of about 35 species distributed 
throughout the arid regions of Central Asia but extending into the Mediterranean Basin, China and 
Afghanistan, and known to feed on legumes within the papilionoid tribes Galegeae, Trifoliae, Loteae and 
Sophoreae. 



8 I. D. HODKINSON & D. HOLLIS 

Key to species 

1 Forewing (Fig. 21) with diffuse reddish brown cloud around apex; radular spinules dense, dark 

brown, at low magnification appearing as small distinct spots around the wing margin. Cell 
mx+2 long, at least as long as the stem of vein M. Vein Cu Xh very strongly curved before its 
junction with the marginal vein. On Glycyrrhiza species glycyrrhizae (Becker) (p. 8) 

- Forewing (Figs 26, 31, 37) without pattern, radular spinules obvious but not forming distinct 

brown marks. Cell m x + 2 shorter than the stem of vein M. Vein Cw lb less strongly curved 
before junction with the marginal vein. Not on Glycyrrhiza 2 

2 Forewing (Fig. 26) with surface spinules occupying all the cells. Male paramere as in Figs 28, 29. 

On Anthyllis pro/iaskaiPreisner(p.9) 

- Forewing (Figs 31 , 37) without surface spinules except in the radular areas. Male paramere as in 

Figs34,40.Noton,4^/iy//w 3 

3 Male paramere (Figs 33, 34) relatively broad, with an anterior subapical bulge. Turkey, on 

Astragalus stocklosai (Klimaszewski & Lodos) (p. 10) 

- Male paramere (Fig. 40) narrower, without an anterior subapical bulge. South-western 

U.S.S.R. , on Medicago and Vicia species medicaginis (Andrianova) (p. 9) 



Cyamophila glycyrrhizae (Becker) 
(Figs 21-25, 93) 

Psyllodes glycyrrhizae Becker, 1864: 486. Syntypes C\ $, U.S.S.R. [not examined]. 

Psylla glycyrrhizae (Becker) Low, 1881: 262; 1883: 239; Puton, 1886: 91; Horvath, 1904: 579; Oshanin, 
1907: 351; 1912: 127; Aulmann, 1913: 16; Sulc, 1910a: 21; Vondracek, 1953: 174; Klimaszewski, 1963: 
427; 1968a: 230; 1969/?: 222; 1973: 208; Loginova, 1964a: 465; 1968: 305; 19726: 140; 1981: 26; Loginova 
& Baeva, 1972: 6; Baeva & Kankina, 1971: 84; Gegechkori, 1969a: 222; 19696: 722; 1978: 20; Halperin et 
a/., 1982: 32. 

Cacopsylla glycyrrhizae (Becker) Klimaszewski & Lodos, 1977: 5. 

Cyamophila glycyrrhizae (Becker) Loginova, 1977: 582; 1981: 26; Klimaszewski & Lodos, 1979: 8; 
Gegechkori, 1981: 119; 19836: 148; 1984: 104; Baeva, 1985: 206. 

Cyamophila odontopyx Loginova, 1978: 86. Holotype cf , U.S.S.R. [not examined]. Syn. n. 

Description. Coloration. General coloration of head and thorax orange-red with white markings, 
abdomen and terminalia somewhat paler; forewing pale yellow with diffuse apical reddish brown pattern 
and dark brown spots in the radular areas; antennal segments 1-8 yellow, segments 3-8 apically darkened, 
segments 9-10 dark brown. All the specimens to hand are immature and it is likely that fully mature 
specimens are darker. 

Structure. Head (Fig. 22) robust, genal cones short and broad, with broadly-rounded apices. Forewing 
(Fig. 21) with well-developed pterostigma and costal break; spinules confined to radular areas where they 
form distinct black marginal punctuations; cell m x elongate, cell cw la high, vein Cu ib strongly recurved 
adjacent to wing margin. Metatibia with 5 thick black spurs, basal metatarsus with 2 black spurs. Male 
proctiger (Fig. 23) simple; paramere (Fig. 24) parallel-sided with a large inner apical tooth; aedeagus (Fig. 
25) with broad hooked apex. Female terminalia (Fig. 93) moderately long. 

Measurements. See Table 1. 

Material examined 

14 0", 6 $ , Israel and U.S.S.R. (IDH); 1 cf , 1 $ , Iran (BMNH). 

Host plant. Glycyrrhiza glabra L. 

Published distribution. Turkey (Klimaszewski & Lodos, 1977, 1979; Vondracek, 1953), Israel (Halperin 
etai, 1982), Afghanistan (Loginova, 19726; Loginova & Baeva, 1972), Iran (Loginova, 19726; Loginova 
& Baeva, 1972), Mongolia (Klimaszewski, 19686), U.S.S.R.: S. European part, Ukrainian SSR, Arme- 
nian SSR, Georgian SSR, Gruz SSR, Azerbaijan SSR, Transcaucasus' , 'Pamir', Kazakh SSR, Uzbek 
SSR, Turkmen SSR, Tadjhik SSR (Gegechkori, 1984; Baeva, 1985). 

Comments. Cyamophila odontopyx falls within the range of variation of C. glycyrrhizae. We are unable to 
separate material of both nominal species, determined by Loginova, and the two are here synonymized. C. 
glycyrrhizae is separated from other west Palaearctic members of the genus by the presence of a subapical 
band of infuscation in the forewing. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 9 

Cyamophila medicaginis ( Andrianova) 
(Figs 37-42) 

Psylla medicaginis Andrianova, 1952: 270; Loginova, 1968: 306; 19726: 141; Gegechkori, 1968: 510; 
19696: 723; 1970: 714; 1976: 65; Baeva & Kankina, 1971: 84; Klimaszewski, 1963: 428; 1969a: 47; 1973: 
214; Gegechkori & Djibladze, 1976: 33. Syntypes, Q\ $ , U.S.S.R. [not examined]. 

Cacopsylla medicaginis (Andrianova) Klimaszewski, 1975: 199. 

Cyamophila medicaginis (Andrianova) Loginova, 1977: 582; 1981: 26; Gegechkori, 1978: 21; 1981: 121; 
1984: 148. 

Description. Coloration. Specimens to hand had the body, legs and antennae orange-yellow throughout. 
Slightly darker markings were present on dorsum of thorax. 

Structure. Forewing (Fig. 37) oblong oval, with spinules confined to radular areas; pterostigma and 
costal break well-developed. Male proctiger (Fig. 39) simple; paramere (Fig. 40) parallel-sided, expanded 
apically into forward pointing tooth; apex of aedeagus slightly reniform. Female terminalia as in Fig. 42. 

Measurements. See Table 1. 

Material examined 

2 d\ 2 ?, U.S.S.R.: Tabagatan, Semipal 6.vii.l962, {Loginova). 

Host plant. Medicago sativa L. and possibly Vicia sp. 

Published distribution. Southern U.S.S.R., Transcaucasus' (Gegechkori, 1968; 1969a; 1970; 1976; 
1978; 1981; 19836; 1984; Gegechkori & Djibladze, 1976), Ukrainian SSR (Klimaszewski, 1963), Pamir 
(Baeva & Kankina, 1971), Kazakh SSR (Andrianova, 1952; Loginova, 19726), Central Asia (Gegechkori, 
1984). 

Comments. This species has yet to be recorded from outside the U.S.S.R. but Klimaszewski (1969, 1975) 
includes it in his key to the Polish fauna on the grounds that as it occurs in the west U.S.S.R. it is likely to 
occur in Poland. We have followed Klimaszewski. It can be separated from the other clearwing west 
Palaearctic Cyamophila species by the following combination of characters: the lack of forewing spinules 
throughout the membrane, the shape of cells cu la and ra x and the shape of the male paramere and 
aedeagus. 

Cyamophila prohaskai (Preisner) 
(Figs 26-30, 94) 

Psylla prohaskai Preisner, 1927: 263; Prohaska, 1928: 4; Haupt, 1935: 234; Franz, 1943: 382; Schaefer, 
1949a: 53; 19496: 31; Wagner & Franz, 1961: 167; Kuwayama & Miyatake, 1971: 52; Klimaszewski, 
1973: 219. Syntypes cf $, Austria [not examined]. 

Cyamophila prohaskai (Preisner) Loginova, 1977: 582; Burckhardt, 1983: 54; Conci &Tamanini, 1986: 60. 

Description. Coloration. General body colour of mature specimens orange-red with white markings on 
dorsum of thorax; forewing membrane clear to pale yellow, veins yellow; antennal segments 1-8 yellow, 
segments 3-8 infuscate apically, segments 9-10 dark brown; legs orange-yellow. 

Structure. Head (Fig. 27) with relatively long genal cones. Forewing (Fig. 26) oblong-oval, with 
well-developed pterostigma and costal break; spinules present throughout all cells. Metatibia with 5 thick 
black apical spurs; basal metatarsus with 2 spurs. Male proctiger (Fig. 28) simple, paramere (Fig. 29) 
gradually broadening to a truncate apex, with an anteriorly directed apical tooth; aedeagus (Fig. 30) 
slender, with a slightly hooked apex. Female terminalia (Fig. 94) long and slender, with dorsal margin of 
proctiger sinuous; circumanal pore ring small. 

Measurements. See Table 1. 

Material examined 
2 cf, 2 $, Switzerland (DB). 

Host plant. Anthyllis vulneraria L. 

Published distribution. Austria (Preisner, 1927; Prohaska, 1928; Haupt, 1935; Franz, 1943; Wagner & 
Franz, 1961), Italy (Conci & Tamanini, 1986), Switzerland (Schaefer, 1949a, 6; Burckhardt, 1983). 

Comments. The record for China published by Kuwayama & Miyatake (1971) is unlikely to be correct: all 
other records are from the European Alps. C. prohaskai can be distinguished from the other west 
Palaearctic clearwing Cyamophila species by the presence of dense spinules on the forewing membrane. 



10 



I. D. HODKINSON & D. HOLLIS 

Cyamophila stoklosai Klimaszewski & Lodos 



(Figs 31-36) 

Cyamophila stoklosai Klimaszewski & Lodos, 1979: 9. Holotype cf, Turkey (Lodos, Klimaszewski 
collection) [not examined]. 

Material of this species was unavailable and the following description is based entirely on information given 
in the original description. The drawings are derived from photomicrographs presented by Klimaszewski & 
Lodos (1979). 

Description. Head and body sandy yellow with rust-coloured patches and bands; abdominal sclerites 
brown, intersegmental membranes yellow; genitalia slightly paler brown than rest of abdomen; forewing 
transparent, slightly yellowish towards apex; antennae yellow except for segments 8-10 and the apices of 
segments 6-7 which are brown; legs yellow with brownish patches on femur. 

Structure. Head (Fig. 32) robust, genal cones short, with broadly rounded apices. Forewing (Fig. 31) 
oblong-oval, with well-developed pterostigma and costal break; spinules confined to the radular areas; cell 
mi narrow; cell cw la high, strongly arched. Male proctiger (Fig. 33) simple; paramere (Fig. 35) with bulbous 
apex. Female terminalia as in Fig. 36. 

Measurements. See Table 1. 

Host plant. Astragalus sp. 

Published distribution. Turkey (Klimaszewski & Lodos, 1979). 



Table 1 Measurements of Acizzia and Cyamophila species. Males and females are given separately and 
all values are in mm. 



Female Male 

Head Antennal Forewing proctiger proctiger 
width length length length length 



Male Male 

paramere aedeagus 

length length 



Acizzia 


















bicolorata 


m 
f 
m 


0-54 


0-93 


1-50 


— 


0-18 


0-18 


0-16 


hollisi 


0-57-0-65 


1-00-1-14 


1-63-1-83 





0-20-0-23 


0-17-0-24 


0-15-0-19 




f 


0-61-0-65 


1-05-1-22 


1-92-2-12 


0-55-0-88 


— 


— 


— 


uncatoides 


m 


0-59-0-63 


0-79-0-96 


1-67-1-80 


— 


0-28-0-29 


0-21-0-22 


0-20-0-21 




f 


0-58-0-61 


0-83-0-88 


1-80-1-91 


0-35-0-39 


— 


— 


— 


acaciae- 


m 


0-34-0-42 


0-55-0-60 


1-17-1-24 


— 


0-12-0-14 


0-12-0-13 


0-12-0-13 


baileyanae 


f 


0-37-0-44 


0-60-0-64 


1-30-1-61 


0-30-0-31 


— 


— 


— 


Cyamophila 


















glycyrrhizae 


m 


0-78-0-80 


0-90-0-96 


2-20-2-31 


— 


0-33-0-35 


0-29-0-31 


0-28-0-29 




f 


0-79-0-81 


1-00-1-10 


2-30-2-32 


0-70-0-72 


— 


— 


— 


medicaginis 


m 
f 
m 






not available 








prohaskai 


0-75-0-78 


1-07-1-10 


2-42-2-44 





0-31-0-33 


0-26-0-27 


0-27-0-29 




f 


0-81-0-83 


1-13-1-16 


2-63-2-64 


0-86-0-89 


— 


— 


— 


stoklosai 


m 


0-62 


0-98 


1-68 


— 


0-32 


0-24 


— 




f 


0-66 


1-10 


1-98 


0-66 


— 


— 


— 



ARYTAINA Forster 

Arytaina Forster, 1848: 69; Pflugfelder, 1941: 76. Type-species: Psylla spartii Hartig, 1841 (= Psylla 

genistae Latreille, 1804), designated by Oshanin, 1912: 128. 
Amblyrhina Low, 1879: 599; 18886: 382; Kieffer, 1905: 165; Heslop-Harrison, 1951: 425; Loginova, 1976a: 

598; Hodkinson & White, 1979a: 59. Type-species: Psylla torifrons Flor, 1861, by monotypy. Syn. n. 
Psyllopa Crawford, 1911: 628. Type-species: Psyllopa magna Crawford, by original designation. [Synony- 

mised by Crawford, 1914: 122.] 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 11 

Diagnosis. Head weakly deflexed, genal cones short and broad, with broadly rounded apices, in 
approximately same plane as vertex. Pronotum broad and flat, propleurites quadrate, divided by a vertical 
suture. Forewing oval to elongate-oval, usually broadest at or before middle; pterostigma and costal break 
absent; cell cu Xa low and rarely slanting towards base of wing. Basal metatarsus with one black spur. Male 
proctiger without posterior lobe; paramere broad with truncate apex, terminated by a characteristic 
anteriorly directed tooth; aedeagus with a strongly hooked apex. Associated with legumes of the tribe 
Genisteae. 

Comments. Arytaina is a monophyletic genus of 11 species associated with host-plants of the legume tribe 
Genisteae within the Mediterranean Basin. It contains the original core species related to the type-species 
plus most species hitherto referred to Amblyrhina. The synonymy given relates only to references not 
quoted under the individual species synonymies and to papers that deal specifically with the generic 
synonymy. Unfortunately Arytaina had become a holding genus for many other species, outside the area 
under study, which are unrelated to the type-species and it is necessary to reallocate these species, where 
possible, to other genera (see Hodkinson, 1983; Hodkinson & White, 1981; Jensen, 1957b; Tuthill, 
1943; Mathur, 1975; Capener, 1970). The following species are here transferred to other genera as 
follows. 



Acizzia obscura (Crawford, 1912; from Psyllopa). Comb. n. 

= A. spinosa (Mathur, 1975; from Arytaina) . Syn. n., Comb. n. 
A. albizziae (Yang, 1984; horn Arytaina). Comb. n. 
Arytainilla devia (Loginova, 1976; fi om Arytaina) . Comb. n. (see p. 42). 
A. nubivaga (Loginova, 1916b; from Arytaina). Comb. n. (see p. 42). 
Ceanothia assimilis (Crawford, 1914; from Arytaina) . Comb. n. 
C. bicolor (Jensen, 1957b; horn Arytaina). Comb. n. 
C. boharti (Jensen, 1957b; from Arytaina). Comb. n. 
C. essigi (Jensen, 1957b; from Arytaina). Comb. n. 
C. insolita (Tuthill, 1943; ir om Arytaina). Comb. n. 
C. mitella (Jensen, 1957b; from Arytaina) . Comb. n. 
Euphalerus isitis (Cotes, 1893; from Psylla). Comb. n. 
E. punctinervis (Crawford, 1919; from Arytaina). Comb. n. 
Euryconus pulchra (Crawford, 1919; from Arytaina). Comb. n. 
Insnesia brevigena (Crawford, 1919; from Arytaina). Comb. n. 
/. crawfordi nom. n. 

= /. pulchra (Crawford, 1920; from Arytaina). Comb. n. (homonym of Arytaina pulchra Crawford, 

1919). 
I.flava (Crawford, 1919; from Arytaina). Comb. n. 
/. iolani (Crawford, 1919; from Arytaina). Comb. n. 
/. meridionalis (Crawford, 1919; from A rytaina). Comb. n. 
/. thakrei (Mathur, 1973; from Arytaina). Comb. n. 
/. tuberculata (Crawford, 1917; from Arytaina). Comb. n. 
/. uichancoi (Braza & Calilung, 1981; from Arytaina). Comb. n. 
/. variabilis (Crawford, 1917; from Arytaina). Comb. n. 
Peripsyllopsis ramakrishni (Crawford, 1912). Stat, rev., Comb. rev. 
Retroacizzia mopanei (Pettey, 1924; from Arytaina). comb. n. 

= R. antennata Heslop-Harrison, 1961b. Syn. n. 
Spanioneura turkiana (Klimaszewski & Lodos, 1977; from Amblyrhina) . Comb. n. 
S. pechai (Klimaszewski & Lodos, 1977; from Amblyrhina). Comb. n. 

Arytaina virgina Caldwell, 1944 and A. cornicola Frauenfeld, 1896 are both regarded here as nomina 
dubia, while A. fasciata Laing, 1930 remains the sole species which cannot be assigned to another genus at 
present. 

Key to species 

1 Forewing (Fig. 43) with cell m 1+2 elongate, as long as stem of vein M. Cell cw la narrowly 

triangular, veins Cw la and Cu lb almost straight longicella sp. n. (p. 16) 

- Forewing (Figs 45, 51, 57) with cell m 1+2 much shorter than length of vein M. Cell cw la not 

narrowly triangular, veins C« la and Cw lb more distinctly curved 2 



12 I. D. HODKINSON & D. HOLLIS 

2 Cells of forewing with distinct longitudinal brown or black pattern extending backwards from 

the apex (Figs 45, 47, 49) 3 

- Cells of forewing without distinct longitudinal patterning (in atlasiensis, adenocarpi and 

angustatipennis (Figs 53, 61, 51) there is occasionally a faint clouding of the wing membrane 

and in the former this is accompanied by the presence of diffuse spots in the apical cells) 5 

3 Forewing (Fig. 45) with costal margin comparatively straight, surface spinules present through- 

out cell r\. Genal cones (Fig. 46) broad with rounded apex. Male paramere (Fig. 66) 
comparatively short and broad. Aedeagus as in Fig. 67 genistae (Latreille) (p. 14) 

- Forewing (Figs 47, 49) with costal margin more strongly curved; surface spinules absent 

throughout most of cell r x . Genal cones either broad with an angular apex (Fig. 48) or more 
slender with rounded apex (Fig. 50). Paramere relatively longer and narrower (Fig. 69, 72). 
Aedeagus as in Figs 70, 73 4 

4 Forewing (Fig. 49) with surface spinules absent from cell c+sc and the base of r l5 and confined 

to narrow bands in the centres of cells r s and m. Genal cones (Fig. 50) slender, with rounded 
apex. Male paramere and aedeagus as in Figs 71-73. Spain and Sicily 

maculate (Low) (Spanish material) (p. 16) 

- Forewing (Fig. 47) with spinules present throughout cells c+sc and r 1 and broadly distributed 

throughout cells r s and m. Genal cones (Fig. 48) broader and with a more angular apex. Male 
paramere and aedeagus as in Figs 68-70. Eastern Europe 

maculate (Low) (Hungarian material) (p. 16) 

5 Forewing (Fig. 51) strongly narrowed to an acute apex. Female proctiger (Fig. 101) long and 

slender, with a relatively small circumanal pore ring angustatipennis (Loginova) (p. 14) 

- Forewing (Figs 55, 59, 61) less strongly narrowed, apex more broadly rounded. Female 

proctiger (Figs 92, 99, 100) less slender, relatively shorter or more robust, with a relatively 
longer circumanal pore ring 6 

6 Forewing shaped as in Fig. 53 with cell c+sc somewhat bowed outwards, vein Cw lb strongly 

curved before its junction with the anal margin; nebulous brown spots present on membrane 
around the apex. (N.B. Several other species have dark spots where the longitudinal veins 
meet the marginal vein.) atlasiensis sp. n. (p. 14) 

- Forewing (Figs 55, 59, 61) of different shape, without nebulous brown spots on membrane 

around apex (if pattern present as in A. torifrons (Fig. 57) then never consisting of spots); 

vein Cu ib less strongly curved before its junction with the marginal vein 7 

7 Forewing (Figs 55, 57) opaque amber or yellow-brown throughout or with suffused brown 

markings along veins around apex 8 

- Forewing (Figs 59, 61) not amber or yellow-brown, and without brown patterning along vein 

margins and around apex 9 

8 Forewing (Fig. 55) elongate oval, amber or yellow-brown throughout. Genal cones (Fig. 56) 

short with a broad base and a very broadly rounded apex. Male paramere (Fig. 74) less 
broad . Apex of aedeagus (Fig . 81) more strongly hooked putonii (Low) (p . 17) 

- Forewing (Fig. 57) more broadly oval, shorter, not amber or yellow-brown throughout; pattern 

somewhat variable but usually consisting of suffused brown markings along vein margins and 
around apex. Genal cones (Fig. 58) slightly longer and less broadly rounded at apex. Male 
paramere (Fig. 83) broader. Apex of aedeagus (Fig. 80) less strongly hooked 

torifrons (Flor) (p. 18) 

9 Forewing (Fig. 59) broadest in basal third, with a narrowly rounded apex; vein Cu la weakly 

arched. Male paramere (Fig. 83) with a subapical lobe on the anterior margin. Aedeagus 

(Fig. 84) with a weakly curved tip hispanica sp. n. (p. 16) 

- Forewing (Figs 61, 257) broadest towards the centre, with a more broadly rounded apex and 

with vein Cu ia more strongly arched. Male paramere (Figs 74, 86, 89) without a subapical 

lobe on the anterior margin . Aedeagus (Figs 87 , 90) usually with a strongly recurved tip 10 

10 Male paramere (Fig. 74) almost as long as proctiger, with at most a short anteriorly directed 

apical tooth. Genal cones (Fig. 258) apparently much shorter. (Figures and description taken 
from the original, all type material is lost.) Libya africana Heslop-Harrison (p. 13) 

- Male paramere (Figs 86, 89) distinctly shorter than the length of the proctiger, with a large 

apical tooth. Genal cones (Figs 62, 64) longer, with broadly rounded apices. France, 
Portugal and Spain 11 

11 Forewing (Fig. 63) with dense spinules broadly distributed throughout all cells. Head (Fig. 64) 

more robust, vertex relatively long. Male paramere (Fig. 89) with a distinct anterior bulge at 
mid length and with a sinuously truncated apex which is developed anteriorly into a massive 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 13 

tooth. Male aedeagus as in Fig. 90. Antennae more than twice as long as head width 

magnidentata sp. n. (p. 17) 
- Forewing (Fig. 61) with less dense spinules which are confined to the apical areas of cells r x , r s , 
ra, m 1+ 2> c M ia> the base of c+sc and around vein Cu 2 . Head (Fig. 62) less robust, vertex 
relatively shorter. Male paramere (Fig. 86) more regularly shaped, curved posteriorly and 
with a smoothly convex apex which is developed anteriorly into a large but shorter tooth. 
Male aedeagus as in Fig. 87. Antennae less than twice head width adenocarpi Low (p. 13) 

Arytaina adenocarpi Low 

(Figs 61, 62, 85-87, 99) 

Arytaina adenocarpi Low, 1880: 552; 1883: 230; 1885: 150; Puton, 1886: 92; Aulman 1913: 32; Fuente, 
1920: 321; Heslop-Harrison, 1951: 434; Ramirez Gomez, 1956: 75; Klimaszewski, 1973: 191; Loginova, 
1916a: 598; Conci & Tamanini, 1984b: 262. Lectotype cf, France, here designated (NM). 

Arytaena adenocarpi Low; Oshanin, 1907: 366; 1912: 128. 

Description. Coloration. All the available specimens are immature. General coloration of head, thorax 
and abdomen and appendages bright green, occasionally orange-yellow; often with orange markings on 
dorsum of head, thorax and abdomen; forewing clear, occasionally with faint brownish clouds; veins 
yellow to brown, always much darker at the points where they meet the marginal vein. 

Structure. Head (Fig. 62) with genal cones swollen, with rounded apices. Forewing (Fig. 61) elongate 
oval; spinules sparse and confined to wing apex, the base of cell c+sc and around vein Cu 2 ; pterostigma and 
costal break absent; cell ra x of normal size, vein Cw la strongly curved. Metatibia with 5 thick black spurs; 
metatarsus with 1 black spur. Male proctiger (Fig. 85) simple; paramere (Fig. 86) parallel-sided, curved 
posteriorly in apical half, with a large inner apical tooth projecting anteriorly; aedeagus (Fig. 87) with 
moderately strongly hooked apex. Female terminalia as in Fig. 99. 

Measurements. See Table 2. 

Material examined 

Lectotype cf, France: 'Gallia, Landes. Type v. Aryt. adenocarpi Low' (NM). Paralectotype $, same 
data as lectotype. 7 cf, 6 $, Portugal and France: 'Haute Pyrenees' (BMNH). In addition there is an 
abundance of material in the BMNH, presumably collected by Heslop-Harrison in Spain or Portugal, 
which does not bear data labels. 

Host plants. Adenocarpus complicatus (L.) Gay and Adenocarpus hispanicus (Lam.) D.C. in Lam. & 
D.C. 

Published distribution. France (Low, 1880; Aulmann, 1913), Spain (Fuente, 1920; Ramirez Gomez, 
1956), Italy (Conci & Tamanini, 19846). 

Comments. This species is closest to A. magnidentata. It can be separated by the form of the forewing 
spinule pattern and the shape of the male paramere and aedeagus. 

Arytaina africana Heslop-Harrison 

(Figs 74, 257, 258) 

Arytaina africana Heslop-Harrison, 1951: 438; Loginova, 1972a: 16; 1976a; 598. Syntypes cf and $ , Libya 
[not traced] . 

Description. Coloration. According to Heslop-Harrison this is a 'greenish or yellowish-brown insect with 
darker brown markings on the head and thorax, and brown maculations on the forewings; strongly 
resembling A. genistae in general appearance'. 

Structure. Head (Fig. 258) with short genal cones. Forewing oval, cells m x and m 2 of moderate size, vein 
Cw la strongly arched; spinules present throughout all cells; pterostigma and costal break absent. Male 
proctiger (Fig. 74) simple; paramere (Fig. 74) long, parallel-sided, with a truncate apex. Shape of aedeagus 
and of female terminalia unknown. 

Measurements. See Table 2. 

Host plant. 'A yellow flowered broom' (Heslop-Harrison, 1951). 

Published distribution. Libya (Heslop-Harrison, 1951) and possibly Morocco, although the specimens 
identified as this species by Loginova (1972) do not fit the original description very well. 



14 I. D. HODKINSON & D. HOLLIS 

Comments. All type material of this species appears to be lost and we have seen no other authentic 
material. Our account is based entirely on Heslop-Harrison's original description. A. africana appears 
closest to A. genistae but the male paramere is more slender and the genal cones are less massive. 

Arytaina angustatipennis (Loginova) comb. n. 

(Figs 51, 52, 81, 101) 

Amblyrhina angustatipennis Loginova, 1972: 24; Loginova, 1976a: 598. Holotype cf , Morocco (ZMU) 
[examined]. 

Description. Coloration. Dorsal surface of head and thorax brownish yellow with longitudinal reddish 
brown and orange markings; genal cones dirty yellow; abdominal sclerites reddish brown, darker above, 
intersegmental membranes yellow; genitalia yellowish brown; fore wing membrane clear to very pale 
yellow; veins yellow basally, becoming dark brown apically; radular spinules prominent, forming dark 
brown spots at the wing margin; antennae yellow basally, darkening towards the apex; legs brownish 
yellow. 

Structure. Head (Fig. 52) with vertex very short but broad; genal cones short and very broadly rounded. 
Forewing (Fig. 51) broadest in basal third, tapered to a narrow acute apex; spinules confined to the radular 
areas; cells m x and cu la of normal size; vein Cw la strongly curved; pterostigma and costal break absent. 
Metatibia with 5 thick black apical spurs; basal metatarsus with 2 spurs. Male proctiger (Fig. 81) simple; 
paramere (Fig. 81) parallel-sided with a truncate apex. Female terminalia (Fig. 101) long and slender. 

Measurements. See Table 2. 

Material examined 

Holotype cf , Morocco: Atlas Mountains, Reraia, 5-15. vi. 1926 (Lindberg) (ZMU). Paratypes, 1 cf , 1 
$ , same data as holotype. 

Host plant. Unknown. 

Published distribution. Morocco (Loginova, 1972). 

Comment. This species is separated from other members of the genus by the apically tapered forewing. 

Arytaina atlasiensis sp. n. 

(Figs 53, 54, 91) 

Description. Coloration. Dorsal surface of head and thorax dirty yellow, with orange or reddish brown 
markings; genal cones yellow. Abdominal sclerites dark brown to black; intersegmental membranes 
yellow; genitalia dark brown to black; forewing membrane clear to pale yellow, veins concolorous; 
nebulous small brownish spots present towards wing apex; antennal segments 1-4 yellow; remainder dark 
brown; legs orange-yellow. 

Structure. Head (Fig. 54) robust, genal cones short but massive, with broadly rounded apices. Forewing 
(Fig. 53) broadly oval, with cell c+sc bulging outwards; costal break and pterostigma absent; vein R { 
reaching wing margin at or about the centre of the wing; cell m x narrow; cell cw la more strongly arched than 
normal; vein Cw lb strongly recurved as it approaches wing margin; spinules confined to cells in basal half of 
wing. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 spur. Male unknown. Female 
terminalia (Fig. 91) long and robust. 

Measurements. See Table 2. 

Holotype $, Morocco: Middle Atlas, 18.V.1961 (Lawrence) (BMNH). 
Paratypes. 3 $ , same data as holotype. 

Host plant. Unknown. 

Comments. This species appears to be closest to A genistae and A maculata but can be distinguished by the 
characteristic shape and pattern of the forewing. 

Arytaina genistae (Latreille) 

(Figs 45, 46, 65-67, 96) 

Psylla genistae Latreille, 1804: 382; Guerin, 1843: 370; Burmeister, 1843: 139; Bolivar & Chicote, 1879: 
183; Reiber & Puton, 1880: 75. Syntypes cf , ? , France [not traced]. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 15 

Psylla ulicis Curtis, 1835: 565 (22a); Low, 1883: 252. Syntypes [incomplete data], Great Britain (UMM). 

[Synonymised by Low, 1877: 126.] 
Psylla spartii Hartig, 1841: 375; Forster, 1848: 69; Lethierry, 1874: 86; Low, 1877: 126; 1883: 251; Flor, 

1861: 358; Szulczewski, 1927: 199. Type [incomplete data], Germany; Berlin. [Synonymised by Low, 

1877: 126.] 
Arytaina spartii (Hartig) Forster, 1848: 69; Low, 1877: 126; Meyer-Dur, 1871: 404. 
Psylla genistae Fabricius; Lethierry, 1869: 365. [Fabricius reference not traced.] 
Arytaena ulicis (Curtis) Scott, 1876: 529. 
Chermes (Ataenia) genistae (Latreille) Thomson, 1877: 828. 
Chermes genistae (Latreille) Douglas, 1878: 41. 
Arytaena genistae (Latreille) Low, 1879: 597; Scott, 1880: 132; 1882a: 14; 18826: 255; Reuter, 1881: 

162; Edwards, 1896: 250; Oshanin, 1907: 366; 1912: 128; Heslop-Harrison, J. W., 1915: 401; Van der 

Goot, 1912: 284; Britten, 1930: 75; Wahlgren, 1934: 89; Haupt, 1935: 241; Murray, 1936: 138; Schaefer, 

1949a: 54; 19496: 31; Vondracek, 19516: 128; Smreczynski, 1954: 141; Glowacka & Harisanov, 1983: 

64. 
Arytaina genistae (Latreille) Low, 1883: 239; 1884: 149; 1888a: 19; Puton, 1886: 92; Hueber, 1904: 275; 

Heslop-Harrison, 1937: 2; 1951: 428; Lauterer, 1971: 197; Loginova, 1976a: 598. (All other references 

listed under distribution refer to Arytaina genistae unless listed above.) 
Psyllopa magna Crawford, 1911: 628. Syntypes d\ $, U.S.A. [Synonymised by Crawford, 1914: 126.] 
Arylaina genistae (Latreille) Halbert, 1934: 310. [Mis-spelling.] 

Description. Coloration. Immature specimens with body coloration bright green often with orange 
or brown markings on the dorsum. Mature specimens with dorsal surface of head and thorax yellowish 
white with extensive longitudinal brown or orange-brown markings; genal cones yellowish white; 
abdominal sclerites dark brown to black; intersegmental membranes yellow; genitalia dark brown; fore- 
wing membrane clear with brown or sometimes greyish longitudinal pattern; veins brown; antennal 
segments 1-5 brownish yellow, remainder dark brown; legs dirty yellow, usually with femora darkened 
above. 

Structure. Head (Fig. 46) moderately robust, genal cones short and moderately broadly rounded. 
Forewing (Fig. 45) oval, pterostigma and costal break absent; spinules present in all cells; cells m x and cw la 
of average size for genus; vein Cw la strongly curved. Metatarsus with 5 thick black apical spurs; basal 
metatarsus with 1 black spur. Male proctiger (Fig. 65) simple; paramere (Fig. 66) relatively broad, 
parallel-sided with a truncate apex which is developed into an anteriorly directed tooth; aedeagus (Fig. 67) 
with strongly hooked apex. Female terminalia as in Fig. 96. 

Measurements. See Table 2. 

Material examined 

The material is too extensive to list. It includes all the material in the BMNH, together with material in 
the collections of IDH and DB. 

Host plants. Occurs primarily on Cytisus scoparius (L.) Link but is also recorded in the literature from 
Chamaecytisus austriacus (L.) Link, Chamaecytisus heuffelii (Wierzb.) Rothm. and Genista tinctoria L. 
These published records require confirmation as they may reflect the confusion between this species and A . 
maculata. 

Published distribution. Austria (Low, 1888a), Bulgaria (Glowacka & Harisanov, 1983), Czechoslovakia 
(Low, 1888a; Duda, 1892, Sulc, 1905a; Vondracek, 1957; Lauterer, 1977), Denmark (Jacobsen, 1919), 
France (Lethierry, 1869; Reiber & Puton, 1880: Carpentier & Dubois, 1892: Dominique, 1902; Lambertie, 
1910), Great Britain (Scott, 1880; Edwards, 1896; Britten, 1930; Heslop-Harrison, J. W., 1915; Heslop- 
Harrison, 1936a,6,c; Murray, 1936; Ing, 1966; 1971; 1974; Watmough, 1968a,6; Hodkinson, 1976; 1978; 
Hodkinson & White, 19796; White & Hodkinson, 1982), German D.R. (Lauterer, 1966; Emmrich, 1976; 
1978), German F.R. (Forster, 1848), Ireland (Halbert, 1935), Italy (Ferrari, 1888), Netherlands (Van der 
Goot, 1912; Blote, 1926), Poland (Szulczewski, 1927; Smerczynski, 1954; Klimaszewski, 1961; 1967; 
1969a; 1971; 1973; 1975; 1979; Zgardinska, 1976; Glowacka, 1979), Spain (Bolivar & Chicote, 1879; 
Vondracek, 19516), Sweden (Reuter, 1881; Wahlgren, 1934; Ossiannilsson, 1952; 1971), Switzerland 
(Schafer, 1949a,6; Burckhardt, 1983), Rumania (Dobreanu & Manolache, 1962), U.S.A. (introduced) 
(Crawford, 1914; Van Duzee, 1917; Tuthill, 1943; Vondracek, 1953). 

Comments. Early records of this species from Hungary have been shown to relate to A. maculata Low 
(Horvath, 1918). The two species are easily confused and the records given above should be treated with 
care, particularly those for eastern Europe and Italy where their ranges appear to overlap. The two species 
can be distinguished by the shape of the forewing and the form of the male genitalia. 



16 I. D. HODKINSON & D. HOLLIS 

Arytaina hispanica sp. n. 

(Figs 59, 60, 82-84, 92) 

Description. Coloration. Dorsal surface of head and thorax brownish yellow with darker brown longi- 
tudinal markings; abdominal sclerites dark brown above, paler beneath; intersegmental membranes dull 
yellow; genitalia yellow-brown to brown; fore wing membrane very pale yellowish orange; veins con- 
colorous; antennal segments 1-2 yellow; segments 3-4 yellow basally, infuscate apically; segments 5-10 
brown; legs brownish yellow. 

Structure. Head (Fig. 60) with genal cones shorter than vertex and broadly rounded at apex. Forewing 
(Fig. 59) semi-opaque, broadest towards base, tapering to narrowly rounded apex; veins thick; spinules 
dense and present in all cells; cell m x of average size; cell cw la small with vein Cw la weakly curved. Metatibia 
with 5 thick black apical spurs; metatarsus with 1 black spur. Male proctiger (Fig. 82) simple; paramere 
(Fig. 83) with distinct subapical bulge on anterior margin; aedeagus (Fig. 84) weakly hooked at apex. 
Female terminalia as in Fig. 92. 

Measurements. See Table 2. 

Holotype d\ Spain: Sierra Nevada, Loma del Mulhacen, 3200 m, 30.viii.1975 (Barfuss) (ETH). 
Paratypes. 6 $ , same data as holotype. 

Host plant. Undetermined 'Genista' species. 

Comments. This species has the short, oval wing as found in A. torifrons but in other respects resembles 
species such as A. adenocarpi. The form of the male genitalia is characteristic. 

Arytaina I on gi eel I a sp. n. 

(Figs 43, 44, 95) 

Description. Coloration. The type specimen is probably immature. General colour of body and append- 
ages dull dirty yellow throughout (possibly greenish in life). Forewing membrane very pale yellow, slightly 
opaque; veins dull yellow. 

Structure. Head (Fig. 44) with genal cones shorter than vertex, very broad and truncate at the apex. 
Forewing (Fig. 43) oval with narrowly rounded apex; lacking spinules except in radular areas and around 
vein Cu 2 \ cell m x very long and narrow; cell cu la triangular, vein Cw ia straight; pterostigma and costal break 
absent. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Female terminalia as 
in Fig. 95. Male unknown. 

Measurements. See Table 2. 

Holotype $, Spain: Sierra de Guadarrama, vii.1926 (Uvarov) (BMNH). 

Host plant. Unknown. 

Comments. This species is based on a single female but the wing venation is so different it should not be 
confused with other species. 

Arytaina maculata (Low) comb. n. 
(Figs 47-50, 68-73, 97) 

Amblyrhina maculata Low, 1886: 157; 18886: 382; Horvath, 1886: 313; 1918: 58; Puton, 1886: 92; Ragusa, 
1907: 237; Oshanin, 1907: 366; 1912: 128; Aulmann, 1913: 32; Haupt, 1935: 241; Vondracek, 19516: 128 
1957: 180; Klimaszewski, 19696: 44; 1970: 422; 1973: 189; 1975: 129; Loginova, 1964: 464; 1976a: 598 
Lauterer, 1977: 98; Dobreanu & Manolache, 1962: 147; Andrianova & Klimaszewski, 1983: 38 
Glowacka & Harisanov, 1983: 64. Syntypes C?, $ Hungary (not located). 

Amblyfrjrhina maculata Low; Low, 1888a: 19. 

[Arytaina genistae (Latreille), partim; Low, 1888a: 19; Horvath, 1886: 314; 1918: 58. Misidentifications.] 

Amblyrhina maculosa Low; Klimaszewski, 1965. [Mis-spelling.] 

Description. Coloration. Immature specimens with body colour bright green throughout. Mature 
specimens with genal cones, vertex and dorsal part of thorax yellowish white with light brown and orange 
markings; underside of head and thorax yellow with dark brown markings; abdominal sclerites dark brown 
to black, intersegmental membranes orange-yellow; genitalia orange-yellow to dark brown; forewing 
membrane clear with brown or sometimes greyish longitudinal pattern; veins brown; antennal segments 
1-5 yellow, 5-10 dark brown; femora dark brown above, remainder of legs dirty yellow. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 17 

Structure. Head (Figs 48, 50) with genal cones shorter than vertex, broad; more slender in the Spanish 
specimens. Forewing (Figs 47, 49) oval, without pterostigma or costal break; with spinules present 
throughout all cells except for r { , where they are at most confined to the apex; in the Spanish material they 
are also absent from cell c+sc; cells m x and cw la of average size, vein Cw la strongly curved. Metatibia with 5 
thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 68) simple; paramere (Fig. 
69) parallel-sided, truncate, with apical forward-pointing tooth; aedeagus (Fig. 70) strongly hooked. In 
Spanish males the paramere is more slender (Figs 71-73). Female terminalia as in Fig. 97. 

Measurements. See Table 2. 

Material examined 

The only material in the Low collection comes from a different locality to that stated in the original 
description. 1 cf , 1 $, Hungary: Szomotor (det. Low) (NM). 4 Q\ 8 $ Italy and Spain (BMNH, PL). 

Host plants. Chamaecytisus ratisbonensis (Schaeffer) Rothm. and Chamaecytisus borysthenicus (Gruner) 
A. Klaskova. 

Published distribution. Bulgaria (Klimaszewski, 1965, 1970; Glowacka & Harisanov, 1983), Czechoslo- 
vakia (Vondracek, 1957; Lauterer, 1977), Hungary (Low, 1886, 1888a; Horvath, 1886, 1918a; Haupt, 
1935), Italy, Sicily (Ragusa, 1907), Rumania (Dobreanu & Manolache, 1962), U.S.S.R.; central and 
southern European part (Loginova, 1964; Andrianova & Klimaszewski, 1983), Yugoslavia (Horvath, 
1918/?). 

Comments. This species is very close to A. genistae with which it has been confused in the past. It is possible 
that some of the published records for A. genistae refer to this species. The specimens from Sicily have 
slightly more slender genal cones and parameres than Low's material from Austria. Similarly the Spanish 
material listed is included under maculata primarily because of the similarity of the wing pattern. In several 
other respects such as the shape of the genae and parameres it is close to adenocarpi. This whole group 
warrants further detailed investigation. The maculata complex is similar in several respects to A. genistae 
but can be distinguished by the shape of the forewing and the form of the male genitalia. 

Arytaina magnidentata sp. n. 

(Figs 63, 64, 88-90) 

Description. Coloration. Unknown; the type-material is slide mounted. Forewing clear, without pattern 
on membrane but veins darkened at the point where they meet the marginal vein. 

Structure. Head (Fig. 64) robust, genal cones large and broadly rounded at apex, slightly shorter than 
length of vertex. Forewing (Fig. 63) oval, without pterostigma or costal break; dense spinules present 
throughout all cells; cells m x and cw la of average size for genus; vein Cwi a strongly curved. Metatibia with 5 
thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 88) simple; paramere (Fig. 
89) with a massive apical tooth; aedeagus (Fig. 90) with a very strongly hooked apex. Female unknown. 

Measurements. See Table 2. 

Holotype d\ Portugal: Rio Mandego, vii.1953 {Heslop -Harrison) (BMNH). 

Host plant. Unknown. 

Comments. Although based on a single male this species is quite distinct from other Arytaina species. It is 
closest to the other clear elongate-oval winged species, A. adenocarpi, but differs in characters of the 
forewing and the shape of the male paramere and aedeagus. 

Arytaina putonii (Low) comb. n. 
(Figs 55, 56, 75-77, 100) 

Amblyrhina putonii Low, 1888/?: 381; Aulmann, 1913: 32. Syntypes d\ $, France (missing from Low 

collection). 
Amblyrhina putoni Low; Puton, 1899: 113; Oshanin, 1907: 365; 1912: 128; Lambertie, 1910: 96; 

Vondracek, 19516: 128; Klimaszewski, 1973: 190; Loginova, 1976a: 598. 

Description. Coloration. General body coloration bright green with orange markings on the dorsum of the 
head, thorax and abdomen; forewing membrane amber coloured, slightly darker towards apex, veins 
concolorous; antennal segments 1-2 green; segments 3-6 yellowish brown, becoming infuscate towards 
the apex; segments 7-10 brown; legs greenish yellow. 



18 



I. D. HODKINSON & D. HOLLIS 



Structure. Head (Fig. 56) short and broad; genal cones very broad basally and appearing to curve round 
to give broadly truncate apices. Forewing (Fig. 55) short and oval, of somewhat rugose texture, narrowly 
rounded at apex; spinules absent except from radular areas; cells mi and cw la of average size, vein Cw la 
weakly curved to straight; costal break and pterostigma absent. Metatibia with 5 thick black apical spurs; 
basal metatarsus with 1 black spur. Male proctiger (Fig. 75) simple; paramere (Fig. 76) parallel-sided with 
large apical anteriorly directed tooth; aedeagus (Fig. 77) with hooked apex. Female terminalia as in Fig. 
100. 

Measurements. See Table 2. 

Material examined 

10 cT, 7 $ , Portugal and Spain (BMNH). 

Host plant. Recorded by Low (1888) from 'Cytisus spinosus' which, in all probability, is Calicotome 
spinosa (L.) Link. 

Published distribution. France (Low, 18886; Lambertie, 1910; Aulmann, 1913; Vondracek, 19516). 

Comments. This species is closest to A. torifrons but differs in the shape and coloration of the forewing, the 
shape of the genal cones and the form of the male aedeagus and paramere. 



Table 2 Measurements oiArytaina species. Males and females are given separately and all values are in 
mm. 

Female Male Male Male 

Head Antennal Forewing proctiger proctiger paramere aedeagus 

width length length length length length length 

1-38-1-59 1-91-2-19 — 0-39-0-42 0-31-0-32 0-28-0-30 

1-55-1-65 2-14-2-45 0-76-0-85 — — — 

— 2-38 — 

(data from original description) 

2-57 0-25? 0-20? 

1-65-1-70 2-70-2-82 0-62-0-80 — — — 



adenocarpi 


m 
f 


0-77-0-83 
0-80-0-89 


africana 


m 
f 

m 
f 


0-78 


angustatipennis 


0-92-0-98 


atlasiensis 


m 






f 


1-00-1-02 


genistae 


m 
f 


0-90-0-96 
0-95-1-05 


hispanica 


m 
f 


0-85 
0-86-0-90 


longicella 


m 
f 


0-83 


maculata 


m 


0-86 




f 


0-86 


magnidentata 


m 
f 

m 
f 


0-79 


putonii 


0-81-0-86 
0-86-0-93 


torifrons 


m 


0-73-0-75 




f 


0-70-0-83 



unknown 
1-56-1-58 2-54-2-56 0-89-0-97 
2-05-2-20 2-35-2-75 — 

1-90-2-25 2-40-3-00 0-89-0-99 

— 2-01 — 
1-27-1-29 2-32-2-34 0-85-0-86 

unknown 

— 2-34 0-96 
1-68 2-35 — 
1-46 2-35 0-91 
1-61 2-12 

unknown 
1-10-1-14 1-90-2-03 — 

1-15-1-16 2-09-2-20 0-72-0-80 
0-85-0-95 1-55-1-59 
0-85-1-00 1-68-2-02 0-64-0-71 



0-39-0-44 0-37-0-39 0-30-0-32 



0-38 



0-31 



0-31 



0-39 0-33 0-33 

0-37 0-33 0-27 

0-36-0-38 0-31-0-32 0-30-0-31 

0-34-0-36 0-25-0-27 0-26-0-28 



Arytaina torifrons (Flor) comb. n. 

(Figs 57, 58, 78-80, 102) 

Psylla torifrons Flor, 1861: 360. LECTOTYPE cf , France (NM), here designated [examined]. 

[Psylla spartiophila Forster sensu Puton, 1871: 437. Misidentification.] 

Amblyrhina torifrons (Flor) Low, 1879: 600; 18826: 259; 1883: 252; 18886: 382; Puton, 1886: 92; Oshanin, 

1907: 365; 1912: 128; Aulmann, 1913: 32; Klimaszewski, 1973: 190; Loginova, 1976a: 598; Hodkinson & 

White, 1979a: 59. 

Description. Coloration. Immature specimens pale greenish yellow throughout. Mature specimens with 
dorsal surface of head and thorax reddish orange; genal cones yellow; abdomen yellowish brown to dark 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 19 

brown; genitalia brown; forewing clear to pale yellowish, veins pale brown to dark brown, pigment 
spreading from the veins onto immediately adjacent areas of the membrane, apical pattern consisting of 
diffuse brown clouds of irregular form; antennal segments 1-6 brownish yellow; segments 7-10 brown; 
legs brownish yellow. 

Structure. Head (Fig. 58) relatively broad, with genal cones convergent in front of frons. Forewing (Fig. 
57) short and broad, oval; without pterostigma or costal break; spinules confined to radular areas; cells m x 
and cu Xd of normal shape; vein Cu Vd weakly curved. Metatibia with 5 thick black apical spurs; basal 
metatarsus with 1 black spur. Male proctiger (Fig. 78) simple; paramere (Fig. 79) relatively broad and 
appearing swollen along anterior margin, with large apical anteriorly directed tooth; aedeagus (Fig. 80) 
with apex strongly recurved. Female terminalia as in Fig. 102. 

Measurements. See Table 2. 

Material examined 

Lectotype 0\ France: 'Gallia, Marseilles, type v. Aryt. torifrons Flor.' (NM). Paralectotype 2> same 
data as lectotype. 78 0\ $ , France and Spain (BMNH, PL, IDH). 

Host plant. Genista hispanica L. 

Published distribution. France (Flor, 1861; Aulmann, 1913; Hodkinson & White, 1979a). 

Comments. This species is closest to A. putonii but differs in the shape and coloration of the forewing, the 
shape of the genal cones and the form of the male paramere and aedeagus. 



LIVILLA Curtis 

Livilla Curtis, 1836: 625; Low, 1863: 106; Puton, 1876: 288; Kieffer, 1905: 163; Oshanin, 1907: 366; 1912: 

128; Pflugfelder, 1941: 75; Heslop-Harrison, 1951: 421; Loginova, 1976a: 599. Type-species: Livilla 

ulicis Curtis, 1836, by monotypy. 
Floria Low, 1879: 590; Kieffer, 1905: 164; Heslop-Harrison, 1951: 421; Oshanin, 1907: 367; 1912: 128; 

Pflugfelder, 1941: 77; Heslop-Harrison, 1951: 421; Loginova, 1976a: 598; Hodkinson & White, 1979a: 

55. Type-species: Psylla pyrenaea Mink, 1859, designated by Oshanin, 1912: 128 (not by Haupt, 1935: 

241 as indicated by Hodkinson & White, 1979a: 55). Syn. n. 
Alloeoneura Low, 1879: 594; Oshanin, 1907: 368; 1912: 128; Pfleugfelder, 1941: 77; Heslop-Harrison, 

1951: 421; Loginova, 1976a: 599. Type-species: Arytaina radiata Forster, 1848, by monotypy. Syn. n. 
Allaeoneura Low; Puton, 1886: 93; Kieffer, 1905: 163. [Mis-spelling.] 
Floria (Floriella) Ramirez Gomez, 1956: 87. Type-species :Psy lla pyrenaea Mink, 1859, by monotypy. 

[Objective synonym of Floria.] Syn. n. 

Diagnosis. Head weakly deflexed, in same plane as thorax, eyes hemispherical, antennae usually slender, 
at least twice as long as head width; genal cones usually elongate and slender, little deflexed from the plane 
of the vertex. Pronotum broad and flat; propleurites quadrate, divided by a vertical suture. Forewing oval 
to oblong-oval, usually membraneous but short and coriaceous in a few species. Costal break present or 
absent; pterostigma, if present, short; cell cw la not tall and usually not strongly arched. Basal metatarsus 
usually with 1 black spur, occasionally with 2 or 0. Male proctiger simple, without posterior process; 
paramere long and slender, tapered to a hooked apex; aedeagus with apex generally rounded, not strongly 
hooked as in Arytaina. Head and thorax often covered in abundant long setae. Associated with legumes of 
the tribe Genisteae. 

Comments. As defined above the genus Livilla probably constitutes a monophyletic group of 31 species 
living on legume hosts of the Genisteae, primarily within the Mediterranean Basin. The single South 
African species annosa, which Heslop-Harrison (19616) designated type-species of the subgenus Brinc- 
kitia, is excluded from the study. The synonymy given above relates only to references not quoted under 
the individual species synonymies and to papers which deal specifically with generic synonymy. The genus 
now contains species previously included under the old generic names Livilla, Floria and Alloeoneura plus 
one species from Amblyrhina. We could find no characters that warrant the recognition of Floria and 
Alloeoneura. The general body form and genitalia of all the species is very similar and they differ from one 
another primarily in the form of the forewing. We regard the development of short , coriaceous forewings in 
some species merely as a specific adaptation for ensheathing the body to conserve water in a dry 
! environment rather than as a generic synapomorphy. Based on characteristics of the forewing, six loosely 
defined groups of species can be recognised: the ulicis-, horvathi-, burckhardti-, radiata- and spartiisuga- 
groups, with a residual assemblage of heterogeneous species. 



20 I. D. HODKINSON & D. HOLLIS 

Key to species 

1 Forewing (Figs 103, 105, 107) coriaceous and strongly convex, oval, little more than twice as 

long as broad (u/j'cis-group) 2 

- Forewing (Figs 109, 115, 125, 139, 153) membraneous, not strongly convex; usually longer and 

more oblong-oval in shape; if short and/or oval (L. burckhardti (Fig. 165) and L. radiata 

(Fig. 125) ) then with apical brown patterning 4 

2 Genal cones (Fig. 104) massive, longer than the vertex. Forewing (Fig. 103) dark brown 

throughout; cell m 1+2 elongate, longer than the stem of vein M x ; wing membrane with short 
distinct furrows arising from and at right angles to the veins. General body colour shining 
dark brown to black throughout ulicis Curtis (p. 23) 

- Genal cones (Fig. 106, 108) shorter. Forewings (Figs 105, 107) yellowish to amber or brown, 

with a whitish basal patch; cell m 1+2 shorter than stem of vein M; membrane without distinct 
furrows. Body either reddish brown with paler markings or dark brown with 2 large white 
streaks which extend onto the base of the wings 3 

3 Forewing (Fig. 105) brown with broad white basal patch, costal margin weakly curved in basal 

third, cell cw la upright, surface spinules absent from most cells. Genal cones (Fig. 106) with 
very broadly rounded apices. Male genitalia as in Figs 170-172. Iberian Peninsula 

bivittata sp. n. (p. 24) 

- Forewing (Fig. 107) yellow to amber throughout, without white basal patch, costal margin 

strongly curved in basal third, cell c+sc appearing to bulge outwards, cell cw la leaning 
towards base of wing, surface spinules obvious in all cells. Genal cones (Fig. 108) acutely 
tapered. Male genitalia as in Figs 173-175. Austria, Switzerland, N. Italy vicina (Low) (p. 24) 

4 Forewing (Fig. 109) broadest in basal third, acutely tapering towards apex. Genal cones (Fig. 

110) very short, extending little beyond the vertex (burckhardti-group) .... cognata (Low) (p. 25) 

- Forewing (Figs 115, 125, 149, 165) broadest at or beyond the middle, not acutely tapering 

towards apex. Genal cones (Figs 116, 126, 150, 166) much longer 5 

5 Forewing (Fig. 165) with subapical brown chevron pattern, wing apex narrowly rounded. 

Female terminalia (Fig. 270) long and slender. Genal cones shorter than the length of the 
vertex (burckhardti-group) burckhardti sp. n. (p. 25) 

- Forewing pattern, if present (Figs 121, 144, 157), not consisting of brown chevrons, wing apex 

more broadly rounded. Female terminalia (Figs 287-292) more robust. Genal cones (Figs 

116, 122, 126, 132) usually at least as long as vertex 6 

6 Forewing (Fig. 153) clear, without colour pattern or suffused apex 

genistae Ramirez Gomez (p. 38) 

- Forewing (Figs 131 , 144, 161) either with a colour pattern or infuscate apically 7 

7 Forewing (Fig. 125) short and broad, at most 2-1 times as long as broad, with colour pattern as 

shown in Fig. 125; cell r x short and broad. Genal cones (Fig. 126) very long and slender 
(radiate-group) radiata (Forster) (p. 29) 

- Forewing (Figs 117, 121, 123, 127, 161) longer and narrower, more than 2-1 times as long as 

broad (except some male L. poggii (Fig. 142)); pattern not as in Fig. 125; cell r x longer and 
narrower. Genal cones (Figs 118, 122, 124) usually shorter, if long (L. spectabilis (Fig. 162)) 
then more robust 8 

8 Forewing pattern (Figs 111, 113, 115, 117, 155) consisting of longitudinal brown streaks 

(Ziorvaf/ii-group) 9 

- Forewing pattern consisting of brown clouds (Fig. 119), small roundish maculations (Fig. 131), 

infuscation of the wing apex and/or the area adjacent to the anal margin (Figs 159, 163) or a 
combination of these characters (Figs 138, 139) 13 

9 Forewing pattern (Fig. Ill) consisting of narrow longitudinal streaks immediately adjacent to 

the veins in the apical half of the wing. Female circumanal pore ring (Fig. 287) small, at most 

0- 19 times length of proctiger nervosa sp. n. (p. 27) 

- Forewing pattern (Figs 113-115, 117, 155) consisting of an anterior streak which may or may 

not be confined to the apex of vein R s , and a posterior streak extending along the anal 
margin. Female circumanal pore ring (Figs 284, 288, 289, 290) at least 0-21 times length of 
proctiger 10 

10 Forewing (Figs 113, 115) with anterior streak confined to the apex of vein R s , not extending into 

the basal half of the wing 11 

- Forewing (Figs 117, 155) with anterior streak extending across vein R s into the basal half of 

wing 12 

11 Forewing (Fig. 113) with anterior streak consisting of a small spot around the apex of vein R s ; 






W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 21 

costal margin of the wing weakly curved so that apex of cell rj is abruptly truncated by vein 
R s . Genal cones (Fig. 114) strongly divergent, outer margins strongly concave, apex broadly 
rounded. Male proctiger (Fig. 182) somewhat lobed posteriorly. Male aedeagus as in Fig. 
184. N. Africa nigralineata sp. n. (p. 27) 

- Forewing (Fig. 115) with a larger spot or streak around vein R s ; costal margin more strongly 

curved so that apex of cell r x is more narrowly truncated by vein /? s . Genal cones (Fig. 116) 
less divergent, outer margins at most weakly concave, apex narrowly rounded. Male 
proctiger (Fig. 185) less obviously lobed. Male aedeagus as in Fig. 187. E. Europe 

horvathi (Scott) (p. 26) 

12 Forewing (Fig. 117) elongate oblong-oval; anterior streak adjoining the costal margin and 

extending basally to just beyond bifurcation of vein R. Genal cones (Fig. 118) longer and 
slenderer. Male aedeagus (Fig. 190) with a hooked apex. C. Europe 

vittipennella (Reuter)(p. 28) 

- Forewing (Fig. 155) more broadly oval; anterior streak not adjoining the costal margin, leaving 

a long clear area, but extending to base of wing. Genal cones (Fig. 156) shorter and more 
robust. Male aedeagus (Fig. 232) with a bulbous apex. Afghanistan klapperichi sp. n. (p. 28) 

13 Forewing (Figs 119, 121, 123) with the two apical branches of the M vein strongly divergent, 

vein M 1+2 meeting the marginal vein well above the wing apex (radiata-group) 14 

- Forewing (Figs 131, 132, 135, 157) with the branches of vein M not strongly divergent, vein 

M 1+2 meeting the marginal vein at or below the wing apex 16 

14 Forewing pattern (Fig. 123) consisting of small round brown maculae which become confluent 

and form an irregular brown pattern in the apical and posteroapical regions of the wing; 
costal margin strongly curved throughout. Dorsum of thorax uniformly shining chocolate 
brown. Genal cones (Fig. 124) conical. On Genista fasselata syriaca (Low) (p. 31) 

- Forewing (Figs 119, 121) without small brown maculae, pattern consisting primarily of brown 

clouds or bold markings in the apical third of the wing, occasionally with slight infuscation in 
the basal half; costal margin almost straight between the points where veins Ri and R s meet 
the marginal vein. Dorsum of thorax not uniformly shining chocolate brown. Genal cones 
(Figs 120, 122) more elongate. On other hosts 15 

15 Veins in apical third of forewing (Fig. 121) dark brown to black, pattern very dark brown; 

occasionally with the basal half of the wing weakly infuscate. Genal cones (Fig. 122) 
divergent. Basal metatarsus with one thick black spur. Male paramere (Fig. 195) with a 
strongly serrated anterior margin and a massive apical tooth. Male aedeagus (Fig. 196) with 
thin shaft and relatively large apex . Host plant unknown lautereri sp . n . (p . 30) 

- Veins in the apical third of forewing (Fig. 119) not dark brown to black, pattern more 

widespread but less intense, mid brown; basal part of wing not infuscate. Genal cones (Fig. 
120) not divergent. Basal metatarsus without thick black spurs. Male paramere (Fig. 192) 
without strongly serrated anterior margin and with a small apical tooth. Male aedeagus (Fig. 
193) with a broader shaft and a relatively small apex. On Retama sp retamae (Puton) (p. 30) 

16 Forewing (Fig. 161) long and narrow, of a characteristic shape, pattern consisting of an 

irregular brown or orange-brown infuscation of the membrane, primarily in the apical third 
and a darker brown spot around vein Cw lb . Genal cones (Fig. 162) very long, at least 1-3 
times length of vertex. Antennae very long, at least 2-85 times head width. On Spartiwn 
junceum spectabilis (Flor) (p. 37) 

- Forewing (Figs 131, 151, 157, 161) relatively shorter and broader, pattern not as above. Genal 

cones (Figs 132, 152, 158, 164) less than 1-3 times length of vertex. Antennae relatively 
shorter, less than 2-85 times head width. On other hosts 17 

17 Forewing (Fig. 157) with distinct chocolate brown colour pattern distributed around the apex 

of vein R s , the apical half and branches of vein M and around veins Cu x . d and Cw lb 
(spartiisuga-group) pseudoretamae sp. n. (p. 35) 

- Forewing (Figs 131, 144, 145, 159) without distinct chocolate brown patterning in apical third; 

if similarly shaped pattern to pseudoretamae present then much paler, more diffuse and 
overlaid with small round maculations 18 

18 Forewing (Fig. 159) slightly rugose, small maculae absent, yellow-brown to orange-brown 

infuscation occupying a continuous band extending around the apex and along the posterior 
wing margin; remainder of membrane lighter; pigment darker along vein margins giving a 
somewhat furrowed appearance. Male proctiger (Fig. 239) with a slight basal posterior lobe 

adusta (Low) (p. 39) 

- Forewing (Figs 131, 139) usually with small maculae present and broadly distributed through- 



22 I. D. HODKINSON & D. HOLLIS 

out the wing, often overlying the areas of infuscation when present. If maculae apparently 
absent (in some specimens of L. variegata and L. maura they are nebulous and indistinct) 
then infuscation distributed as in Fig. 169, or forewing membrane uniformly yellow to orange 
(Fig. 151). Male proctiger (Figs 203, 206, 209, 245) without basal posterior lobe 19 

19 Forewing (Figs 127, 131, 135, 137, 142) with small brown maculae distributed throughout 

(spartiisuga-group) 20 

- Forewing (Figs 151, 163) with maculae absent or very indistinct; when present very nebulous 

and confined to the apical half of wing 30 

20 Sexually dimorphic species. Male forewing (Fig. 142) with a pale maculation-free band running 

transversely across the centre. Female forewing (Fig. 141) maculate throughout; vein M very 
strongly arched in both sexes, cell cu x very short and high. Apex of male aedeagus as in Fig. 
226. On Genista Corsica in Sardinia poggii (Conci & Tamanini) (p. 36) 

- Forewing (Figs 129, 131, 133, 137) without transverse pale band, occasionally with a pale 

longitudinal band; vein M less strongly arched and cell cw la flatter and more elongate. Apex 

of male aedeagus (Figs 217, 220, 223) of different shape. On other host plants 21 

21 Ground coloration of forewing membrane (Figs 127, 129, 131, 133), excluding maculae, 

uniform , clear to pale yellow throughout 22 

- Ground coloration of forewing membrane (Figs 135, 144, 145, 147, 149), excluding maculae, 

two-tone, clear to pale yellow along anterior half of wing, pale brown to orange-brown along 
posterior half and/or around the apex 27 

22 Maculae absent from a broad band running along anterior margin of forewing (Fig. 133) ; wing 

with a somewhat truncate apex. Male and female genitalia as in Figs 212-214, 274. On 
Calicotome spinosa and Genista pilosa pyrenaea (Mink) (p . 34) 

- Maculae present in area along anterior margin of forewing (Figs 129, 131, 137); wing shaped 

differently, with a less truncate apex. Male and female genitalia as in Figs 206-208, 209-211, 
221-223,271-272. On other hosts 23 

23 Maculae absent from a longitudinal band running the length of cell r s (Fig. 137) . Male paramere 

(Fig. 219) long and curved posteriorly in apical half. Apex of aedeagus (Fig. 220) smoothly 
rounded spartiisuga (Puton) (p. 31) 

- Maculae present throughout cell r s (Figs 127, 129, 131, 139). Male paramere (Figs 204, 207, 

222) usually shorter and broader, if long (Fig. 210) then straighter. Apex of aedeagus (Figs 

205, 208, 211, 223) more angular 24 

24 Forewing (Figs 129, 131) long and parallel-sided, costal margin beyond the pterostigma weakly 

curved to straight ; cell cw la elongate 25 

- Forewing (Figs 127, 139) broader and more oval, costal margin beyond the pterostigma 

strongly curved, cell cu la shorter and more upright 26 

25 Vein R s of forewing (Fig. 129) strongly curved upwards to meet the costal margin, cell r x broad; 

veins M and Cu with a longish common stem. Male paramere (Fig. 207) broad basally, 
gradually narrowing and curving posteriorly towards the apex; basal posterior margin with 
conspicuous thick setae. Male aedeagus (Fig. 208) short and robust. Genal cones (Fig. 130) 
long and slender. Smaller species; head width less than 0-7 mm. Iberian Peninsula and 
N. Africa maculipennis sp. n. (p. 33) 

- Vein R s of forewing (Fig. 131) more sinuous, less strongly curved towards costal margin, cell r x 

narrower; veins M and Cu with a short common stem. Male paramere (Fig. 210) long and 
thin, posterior margin with normal setae. Male aedeagus (Fig. 211) longer and more slender. 
Genal cones (Fig. 132) shorter and more robust. Larger species; head width greater than 0-7 
mm. Sicily siciliensis sp. n. (p. 32) 

26 Forewing (Fig. 127) with a broadly rounded apex; vein R s weakly sinuate, vein Cw la weakly 

curved in apical half. Male paramere (Fig. 204) broad and parallel-sided. Weakly tapered 
towards the apex. Apex of male aedeagus as in Fig. 205. Spain and S. France 

cataloniensis (Hodkinson & White) (p. 32) 

- Forewing (Fig. 139) with a narrowly rounded apex; vein R s strongly sinuate, vein Cw la strongly 

curved in apical half. Male paramere (Fig. 222) narrower and more tapered towards the 
apex. Apex of male aedeagus as in Fig. 223. Greece and Crete hodkinsoni (Burckhardt) (p. 33) 

27 Forewing (Fig. 199) with maculae around apex of vein R s dark brown to black, distinct, much 

darker than those elsewhere on the wing; brown ground colour not extending right round the 
wing apex; wing oval, broadest in the middle, with a narrowly rounded apex. Male paramere 
and aedeagus as in Figs 254-256. Corsica and Sardinia bimaculata sp. n. (p. 34) 

- Forewing (Figs 135, 144, 145, 147) with maculae around apex of vein R s similar in colour to 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 23 

those elsewhere on the wing; brown ground colour extending further around the wing apex; 
wing oblong oval, broadest in apical third, with a broadly rounded apex. Male paramere and 
aedeagus as in Figs 215-217, 236-238, 251-252 28 

28 Larger species, head width greater than 0-70 mm, wing length greater than 21 mm. Male 

paramere (Fig. 216) long and narrow, with a kink at mid length and without very thick basal 
posterior setae. Male aedeagus (Fig. 217) very long and narrow. Genal cones (Fig. 136) 
somewhat divergent. Forewing (Fig. 135) usually without clear pale areas at the apices of 
cells r s , m 1+2 andm. Sicily on Genista aetnensis magna sp. n. (p. 36) 

- Smaller species, head width less than 0-70 mm, wing length less than 2- 10 mm. Male paramere 

(Figs 237, 252) shorter and broader, without a kink at mid length but with conspicuous basal 
posterior setae. Male aedeagus (Figs 238, 253) shorter. Genal cones (Figs 146, 148) less 
divergent. Forewing (Figs 144, 145, 147) often with clear areas at the apices of cells r 8 , ra 1+2 
and m. Not known from Sicily; on other hosts 29 

29 Forewing (Figs 144, 145) relatively narrow, cell r x elongate, cell cw la moderately long, 

patterning as in Fig. 145 although in some specimens, including the type series, the extent of 
the markings is reduced (Fig. 144). Male paramere (Fig. 237) narrowly tapered to apex. Male 
aedeagus (Fig. 238) with apex weakly hooked. Genal cones (Fig. 146) long and slender. On 
Genista hirsuta blandula (Horvath) (p. 35) 

- Forewing (Fig. 147) broader, cell r x shorter and broader, cell cw la shorter and more upright. 

Male paramere (Fig. 252) more broadly tapered to apex. Male aedeagus (Fig. 253) with apex 
strongly hooked. Genal cones (Fig. 148) slightly shorter and more robust. On Genista 
triacanthos lusitanica sp. n. (p. 36) 

30 Forewing (Fig. 163) long, broadest in apical third, veins very fine and indistinct, concolorous 

with membrane; membrane whitish, often with apex infuscate; when infuscate then pale 
areas absent from apices of cells r s , ra 1+2 and m. Genal cones (Fig. 164) slender, longer than 
vertex. Female circumanal pore ring (Fig. 269) less than 0-3 times proctiger length. Larger 
species, head width greater than 0-67 mm, wing length greater than 2-6 mm. Male genitalia as 
in Figs 245-247. Europe, on Laburnum spp variegata (Low) (p. 38) 

- Forewing (Fig. 151) relatively shorter and broader, broadest in middle, veins more robust, 

darker than membrane, apex not infuscate, membrane yellow to pale amber with small paler 
patches at the apex of cells r s , m and m 1+2 . Genal cones (Fig. 152) more robust, shorter than 
the vertex. Female circumanal ring (Fig. 282) relatively longer, more than 0-3 times proctiger 
length. Smaller species, head width less than 0-66 mm, wing length less than 2-5 mm. Male 
unknown. N. Africa, host unknown maura (Vondracek) (p. 39) 

The u/ic/s-group 

The species in this group all possess short, oval, coriaceous forewings. 

Livilla ulicis Curtis 

(Figs 103, 104, 167-169, 263) 

Livilla ulicis Curtis, 1836: 625; Forster, 1848: 68; Meyer-Diir, 1871: 404; Scott, 1876: 528; 1882a: 14; 18826 
255; Reiber & Puton, 1880: 75; Low, 1883: 252; Horvath, 1886: 314; 1918a: 58; Ferrari, 1888: 76 
Edwards, 1896: 250; Dubois, 1898: 239; Puton, 1899: 113; Hueber, 1904: 276; Olivier, 1904: 96 
Oshanin, 1907: 366; 1912: 128; Reuter, 1909: 63; Joakimov, 1909: 1; Lambertie, 1910: 96; Aulmann 
1913: 33; Prohaska, 1928: 5; Haupt, 1935: 231; Schaefer, 1949a: 19; 19496: 30; Vondracek, 19516: 128 
1957: 185; Tamanini, 1955: 11; Ramirez Gomez, 1956: 96; Wagner & Franz, 1961: 163; Dobreanu & 
Manolache, 1962: 146; Lauterer, 1965: 174; 1977: 98; Loginova, 1964: 462; 1976a: 599; Klimaszewski, 
1964: 65; 1965: 203; 1967: 14; 1969a: 43; 1973: 195; 1975: 133; Hodkinson & White, 19796: 39; White & 
Hodkinson, 1982: 26; Burckhardt, 1983: 55. Syntypes cf , $, Great Britain: England (UMM). 
Psylla coleoptera Waltl, 1837: 277. Types (not stated), Germany. [Synonymised by Low, 1883: 252.] 
Psylla callunae Rudow, 1875: 7. Types (not stated), Germany. [Synonymised by Low, 1883: 252.] 

Description. Coloration. Mature specimens with dorsal surface of head and thorax shining, dark brown; 
sclerites of abdomen dark brown, intersegmental membranes reddish yellow; genitalia brown; forewing 
uniform shining brown, veins concolorous; antennal segments 1-3 yellow; segments 4-10 dark brown; legs 
yellowish brown. 

Structure. Head (Fig. 104) with massive sub-cylindrical genal cones, much longer than vertex. Forewing 
(Fig. 103) oval short, coriaceous, strongly convex; costal break and pterostigma absent; spinules, if 



24 I. D. HODKINSON & D. HOLLIS 

present, confined to radular areas; vein R s almost straight; cell m 1+2 elongate; vein M\ reaching to wing 
apex; cell cw la small and flat; vein Cw la almost straight; short furrows running through membrane at right 
angles to the veins. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male 
proctiger (Fig. 167) simple; paramere (Fig. 168) with a few slightly thickened basal setae on the posterior 
margin; aedeagus (Fig. 169) with bulbous apex. Female terminalia as in Fig. 263. 
Measurements. See Table 3. 

Material examined 
This is too extensive to list but includes all the material deposited in the BMNH. 

Host plants. Genista tinctoria L. and possibly Ulex europaeus L. The record for Ononis spinosa L. by 
Ferrari (1888) is unlikely to be correct. 

Published distribution. Austria (Prohaska, 1928; Haupt, 1935); Bulgaria (Joakimov, 1909; Klimaszews- 
ki, 1965), Czechoslovakia (Vondracek, 1957; Lauterer, 1965, 1977; France (Reiber & Puton, 1880; 
Dubois, 1898; Olivier, 1904; Reuter, 1909; Lambertie, 1910), German D.R. (Forster, 1848), German F.R. 
(Haupt, 1935), Great Britain (Curtis, 1836; Scott, 1876; Edwards, 1896; Hodkinson & White, 1979/?; 
White & Hodkinson, 1982), Hungary (Low, 1888a; Horvath, 1886, 1918a), Italy (Ferrari, 1888; Von- 
dracek, 1951/?; Tamanini, 1955), Poland (Klimaszewski, 1964a, 1967, 1969a, 1975), Rumania (Dobreanu 
& Manolache, 1962), Switzerland (Schaefer, 1949a,b; Burckhardt, 1983), U.S.S.R.: European part 
(Loginova, 1964). 

Comments. Livilla ulicis is closest to L. bivittata from which it can be distinguished by the longer genal 
cones and the shape of cells cu x and m x of the forewing. 

Livilla bivittata sp. n. 

(Figs 105, 106, 170-172,264) 

Description. Coloration. Mature specimens with dorsal surface of head and thorax dark shiny brown with 
a pale median streak running medially along head and thorax from base of genal cones to apex of scutellum ; 
further pale streaks running across humeral region and extending on to wing base; abdominal sclerites and 
underside of head and thorax paler yellowish brown; terminalia brown; forewing membrane brown with a 
pale longitudinal streak at the base, veins concolorous; antennal segments 1-2 yellow; segments 3-5 
brownish yellow, apically infuscate; segments 6-10 dark brown; legs brownish yellow. 

Structure. Head (Fig. 106) with short cylindrical genal processes which are evenly rounded at the apex. 
Forewing (Fig. 105) coriaceous, strongly convex; lacking spinules; short, oval shaped; vein R s straight; cell 
mj and cw la small, vein M 1+2 reaching to just below wing apex; pterostigma and costal break absent. 
Metatibia with 5 thick black apical spurs; metatarsus with 1 black spur. Male proctiger (Fig. 170) simple; 
paramere (Fig. 171) without stout basal posterior spines; aedeagus (Fig. 172) with slightly hooked apex. 
Female terminalia as in Fig. 264. 

Measurements. See Table 3. 

Holotype d\ Portugal: Villa Formosana, 1954 (Hes lop -Harris on) (BMNH). 

Paratypes. 1 cf , 1 $ same data as holotype, 3 cf, 2 $ from unknown locality, but intermixed with 
specimens from Spain and Portugal (Heslop-Harrison) (BMNH). In addition there is a lot of material of 
this species without locality labels and in poor condition in the Heslop-Harrison collection (BMNH) which 
is not included in the type series. 

Host plant. Unknown. 

Comments. This species is closest to Livilla ulicis, but can be distinguished from it by the shorter genal 
cones and the shape of cells mj and cu Y of the forewing. 

Livilla vicina (Low) comb. n. 
(Figs 107, 108, 173-175,265) 

Floria vicina Low, 1886: 159; 1888a: 20, Puton, 1886: 92; Hueber, 1904: 276; Oshanin, 1907: 367; 1912: 
128; Aulmann, 1913: 35; Prohaska, 1928: 5; Haupt, 1935: 242; Klimaszewski, 1973: 195; Loginova, 
1976a: 599; Burckhardt, 1983: 55. Syntypes C?, $ , Austria [missing from Low collection, NM]. 

Arytaena montana Cerutti, 1939a: 448; 19396: 583; Schaefer, 1949a: 55; 19496: 31. Lectotype cf, 
Switzerland, designated by Burckhardt, 1983 (MZL). [Synonymised by Burckhardt, 1983: 55.] 

Arytaina montana (Cerutti) Heslop-Harrison, 1951: 435; Tamanini, 1955: 11; 1977: 109; Klimaszewski, 
1973: 191, Loginova, 1976a: 598. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 25 

Description. Coloration. Genal cones, vertex and thoracic dorsum dirty orange-yellow, with indistinct 
darker orange markings on thorax; abdominal sclerites brown, intersegmental membranes yellowish 
orange; genitalia dirty yellow to brown; forewing coriaceous, membrane yellow to orange, veins concolor- 
ous; antenna orange-yellow basally becoming darker towards apex; legs orange-yellow. 

Structure. Head (Fig. 108) with genal cones slightly shorter than vertex, proximate, with rounded apices. 
Forewing (Fig. 107) short and broad, oblong-oval; cell c+sc bulging outwards; costal break and ptero- 
stigma absent; spinules present throughout all cells; vein Rs sinuous, not strongly curved at apex; cell m 1+2 
narrow; vein M 1+2 reaching margin below wing apex; cell cw la large and leaning towards base of wing; vein 
Cu x . d weakly curved. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male 
proctiger (Fig. 173) simple; paramere (Fig. 174) slender, without thickened setae on basal posterior 
margin; aedeagus (Fig. 175) with strongly recurved apex. Female terminalia as in Fig. 265. 

Measurements. See Table 3. 

Material examined 

9 O", 11 $, Switzerland and Italy (BMNH, DB, IDH, MM). 

Host plant. Genista radiata (L.) Scop. 

Published distribution. Austria (Low, 1886, 1888a; Prohaska, 1928; Haupt, 1935), Italy (Tamanini, 
1955, 1977), Switzerland (Cerutti, 1939a,/?; Schaefer, 1949a,6; Burckhardt, 1983). 

Comments. It is a measure of the confusion that has existed within the Arytainini that this species has 
belonged simultaneously to two separate genera. L. vicina resembles L. ulicis and L. bivittata in possessing 
short oval coriaceous fore wings, but it can be distinguished from both these species by the shape and 
detailed venation of the forewing and by the shape of the head and genal cones. 

The burckhardti-group 

The two members of this group have the forewing apex narrowly rounded, and very short genal cones. 

Li villa burckhardt i sp. n. 

(Figs 165, 166, 248-250, 270) 

Description. Coloration. Specimens in the type series appear to have been taken while developing their 
full adult coloration. General body colour (preserved in alcohol) dirty yellow, possibly green in life; 
sclerites of abdomen beginning to turn brown; forewing clear, with brown colour pattern; veins yellow; 
antennae and legs concolorous with body. 

Structure. Head (Fig. 166) with short, broad genal cones. Forewing (Fig. 165) short and oval; 
pterostigma and costal break absent; spinules present throughout all cells; vein Rs strongly curved towards 
marginal vein; vein M evenly curved; vein M x+2 reaching to just below wing apex; vein Cu u strongly 
curved. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 
248) simple; paramere (Fig. 249) relatively short and broad; aedeagus as in Fig. 250. Female terminalia 
(Fig. 270) long and slender with dorsal margin of proctiger shallowly convex. 

Measurements. See Table 3. 

Holotype cf , Spain: Pyrenees, Ordesa National Park, 160 m, on spiny Genista (Lienhard) (ETH). 
Paratypes. 1 cf , 2 $ , same data as holotype. 

Host plant. The type series was collected from a spiny Genista species. 

Comments. L. burckhardti appears to be closest to L. cognata but the resemblance may be superficial. It 
can be separated from all other members of the genus by the chevron patterning in the forewing. 

Li villa cognata (Low) comb. n. 
(Figs 109, 110, 176-178,266) 

Amblyrhina cognata Low, 18826: 258; Oshanin, 1907: 365; 1912: 128; Puton, 1886: 92; Haupt, 1935: 241; 
Vondracek, 19516: 128; 1957: 178; Wagner & Franz, 1961: 162; Lauterer, 1965: 174; 1977: 98; Loginova, 
1976a: 598; Klimaszewski, 1970: 420; 1973: 189; Gegechkori, 1984: 147; Tamanini, 1977: 107; Glowacka 
& Harisanov, 1983: 64. LECTOTYPE cf , Austria, here designated (NM) [examined]. 

Ambly[r]rhina cognata Low; Low 1888a: 19. [Mis-spelling.] 

Description. Coloration. General colour of head, thorax and abdomen bright green, often with orange 



26 ID. HODKINSON & D. HOLLIS 

markings on dorsum of thorax; forewing membrane clear, occasionally with faint brown clouds in apical 
half of wing; veins pale, weakly pigmented; antennae green, apical segments infuscate, legs green. 
Specimens to hand are probably immature and it is likely this species darkens as it becomes older. 

Structure. Head (Fig. 110) short and broad, with small broadly rounded genal cones. Forewing (Fig. 
109) oval, with a narrowly rounded apex; costal break and pterostigma absent; spinules confined to apices 
of cells; vein R s not strongly curved to wing margin; vein Mi evenly curved; vein M 1+2 reaching to just 
below wing apex; cell cu la flat; vein Cw la weakly curved. Metatibia with 5 thick black spurs at apex; basal 
metatarsus with 1 black spur. Male proctiger (Fig. 176) simple; paramere (Fig. 177) slender, without 
thickened basal posterior spines; aedeagus (Fig. 178) with a weakly angled apex. Female terminalia as in 
Fig. 266. 

Measurements. See Table 3. 

Material examined 
Lectotype cf , Austria: 'Austria inf., Modling, det Low' (NM). 
Paralectotype $, same data as lectotype. 2 (J, 2 $ Austria (BMNH). 

Host plants. Chaemaecytisus ratisbonensis (Schaeffer) Rothm., Chamaecytisus supinus (L.) Link, 
Genista germanica L. 

Published distribution. Austria (Low, 1882/?, 1888a; Vondracek, 19516; Wagner & Franz, 1961), 
Bulgaria (Klimaszewski, 1970; Glowacka & Harisanov, 1983), Czechoslovakia (Lauterer, 1965, 1977; 
Vondracek, 1957), France (Aulmann, 1913), 'Germania' (Haupt, 1935), Italy (Tamanini, 1977), 
U.S.S.R.: Caucasus (Gegechkori, 1983). 

Comments. This species has been separated from those with which it had previously been placed within the 
genus Amblyrhina. The characters of the genitalia suggest that it belongs in Livilla, even though the genal 
cones are much shorter than is usual for the genus. It is distinguished further from other Livilla species by 
the narrowly rounded apex of the forewing. 

The horvathi-group 

Members of this group are recognised by the longitudinal brown banding patterns on the forewing 
membrane. 

Livilla horvathi (Scott) comb. n. 

(Figs 115, 116, 185-187,289) 

Floria horvathi Scott, 1879: 84; Low, 1883: 240; 1888a: 20; Horvath, 1886: 314; 1918a: 58; Puton, 1886: 93; 

Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 34; Klimaszewski, 1964: 64; 1967: 14; 19686: 781; 1969a: 

46; 1973: 193; 1975: 130; Loginova, 1964: 464; 1976a: 599; Vondracek, 1957: 192; Lauterer, 1977: 98; 

Klimaszewski & Lodos, 1979: 7; Glowacka & Harisanov, 1983: 64; Onucar, 1983: 56. Holotype ?, 

Hungary [not traced]. 
[Floria vittipennella Reuter sensu Vondracek, 19516: 128; Smrezynski, 1954: 141. Misidentifications.] 

Description. Coloration. Body colour of immature specimens green throughout. Mature specimens with 
vertex and thoracic dorsum dirty greenish yellow with brown markings; genal cones dirty yellow to pale 
brown; abdominal sclerites dark brown, intersegmental membranes greenish yellow; genitalia yellow- 
brown to brown; forewing membrane clear, veins dirty yellow to pale brown; colour pattern orange- 
brown; antennal segments 1-4 dirty yellow, 2-4 apically darkened, remainder brown; legs brownish 
yellow, femora occasionally brown above. 

Structure. Head (Fig. 116) with long divergent genal cones with narrowly rounded apices. Forewing 
(Fig. 115) elongate oval, without pterostigma or costal break; fine spinules distributed throughout all cells; 
vein Rs moderately strongly curved towards margin; vein M slightly sinuous; vein M 1+2 reaching to wing 
apex; cell cw la moderately long but not high; vein Cw la evenly arched. Metatibia with 5 thick black spurs at 
apex; basal metatarsus with 1 black spur. Male proctiger (Fig. 185) simple; paramere (Fig. 186) without 
stout setae on basal posterior margin; aedeagus (Fig. 187) with curved bulbous apex. Female terminalia as 
in Fig. 289. 

Measurements. See Table 3. 

Material examined 

1 CM $, 'Scott collection' [no further data] (BMNH). 16 cf , 16 ?, Greece, Bulgaria and Czechoslo- 
vakia (DB, MM, IDH). 
Host plant. Chamaecytisus austriacus (L.) Link. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 27 

Published distribution. Bulgaria (Klimaszewski, 1967; Glowacka & Harisanov, 1983), Czechoslovakia 
(Vondracek, 1957; Lauterer, 1977), Greece (Vondracek, 1951/?), Hungary (Low, 1888a; Horvath, 1886, 
1918a), Poland (Klimasezewski, 1964, 1967, 19686, 1969a, 1975; Smrezynski, 1954), Rumania (Dobreanu 
& Manolache, 1962), Turkey (Klimasezewski & Lodos, 1979; Onucar, 1983), European U.S.S.R. 
(Loginova, 1964). 

Comments. This appears to be the sister species of L. nigralineata. The two species differ from each other in 
the distribution of brown markings on the forewing and in the shape of the genal cones and male genitalia, 
particularly the aedeagus. 



Li villa nigralineata sp. n. 
(Figs 113, 114, 182-184, 288) 

Description. Coloration. The type series comprises specimens which probably have not developed their 
full adult coloration. General colour pale green throughout with orange markings on the dorsum of the 
head, thorax and abdomen. Forewing membrane clear, wing pattern reddish brown. Antennae with 
segments 6-10 dark brown. 

Structure. Head (Fig. 114) with genal cones widely divergent, longer than vertex, with evenly rounded 
apices. Forewing (Fig. 113) elongate oval, almost parallel-sided; costal break and pterostigma absent; 
spinules fine, present in all cells but restricted in the cells along the anterior edge of wing; cell cw la 
moderately long; vein Cw la moderately strongly arched. Metatibia with 5 thick black apical spurs; basal 
metatarsus with 1 thick black spur. Male proctiger (Fig. 182) simple; paramere (Fig. 183) relatively stout, 
parallel-sided; aedeagus (Fig. 184) with slightly hooked apex. Female terminalia as in Fig. 288. 

Measurements. See Table 3. 

Holotype d\ Algeria: Mtns de Belezma, 1500 m, 5.viii.l980 (Remane) (BMNH). 
Paratypes. 1 cf , 2 $, same data as holotype. 

Host plant. Unknown. 

Comments. This appears to be the sister species to horvathi. It is separated from the latter by the smaller 
patch of brown markings around the apex of vein Rs and by the more divergent genal cones. 



Li villa nervosa sp. n. 

(Figs 111, 112, 179-181,287) 

Description. Coloration. Mature specimens with head, thorax and abdomen pale dirty yellow; dorsum of 
thorax and vertex with pale orange or pale brown markings ; abdomen yellowish or green , turning brownish 
in older specimens; genitalia yellow to brownish yellow; forewing membrane clear to pale yellow, veins 
yellow; patterning pale brown; antennal segments 1-2 yellow, segments 3 to 5 darkened apically, segments 
6 to 10 dark brown; legs yellow. 

Structure. Head (Fig. 112) with slender divergent genal cones, slightly longer than vertex. Forewing 
(Fig. Ill) elongate oblong-oval, almost parallel-sided; costal break and pterostigma absent; fine spinules 
present throughout all cells. Vein Rs strongly curved towards wing margin; vein M long and evenly curved; 
vein M 1+2 meeting margin well above wing apex; cell cw la moderately long. Metatibia with 5 thick black 
apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 179) simple; paramere (Fig. 180) 
curved posteriorly, with thickened setae present on basal posterior margin; aedeagus (Fig. 181) with 
angular bulbous apex. Female terminalia (Fig. 287) with very short circumanal pore ring. 

Measurements. See Table 3. 

Holotype Q\ Spain: Cordoba, Embalse de Iznajar, nr Iznajar, 12. vi. 1977, {Hollis) (BMNH). 
Paratypes. Spain: 8 d\ 15 $, same data as holotype; 1 $, 1 nymph, Andalusia, Porto de los Alazores, 
800 m, 30.viii.1975 {Lienhard) (DB). 

Host plant. Genista radiata (L.) Scop. 

Comments. The upturned vein M 1+2 of the forewing also resembles that found in members of the 
radiata-groxxp; otherwise L. nervosa resembles members of the horvathi-group. It is distinguished by the 
characteristic forewing pattern. 



28 I. D. HODKINSON & D. HOLLIS 

Li villa vittipennella (Reuter) comb. rev. 
(Figs 117, 118, 188-190, 290) 

Psylla vittipennella Reuter, 1875: 333. Syntypes cf , $ Yugoslavia [not located]. 

Floria vittipennella (Reuter) Low, 1879: 593; 1888a: 20; Puton, 1886: 93; Hueber, 1904: 276; Oshanin, 
1907: 367; 1912: 128; Prohaska, 1928: 5; Haupt, 1935: 242; Burckhardt, 1983: 54. 

Livilla vittipennella (Reuter) Aulmann, 1913: 34. 

Floria lineata Cerutti, 1939a: 445; 19396: 583; Schaefer, 1949a: 55; 19496: 31; Klimaszewski, 1973: 193; 
Loginova, 1976a: 599; Tamanini, 1977: 111. Lectotype cf , Switzerland (MZL) designated by Burc- 
khardt, 1983. [synonymised by Burckhardt, 1983: 54.] 

Floria vitipennella (Reuter); Klimaszewski, 1973: 195; Loginova, 1976a: 599. 

Description. Coloration. Mature specimens with dorsum of thorax and vertex brownish yellow with 
orange markings; genal cones yellow; remainder of body orange-yellow; forewing membrane clear, veins 
yellow; colour pattern reddish brown; antennal segments 1 and 2 yellow-orange, remainder pale brownish 
becoming darker towards antennal apex; legs orange-yellow. In some less mature specimens the body 
coloration often has a greenish tinge. 

Structure. Head (Fig. 118) with genal cones long and slender and with narrowly rounded apices. 
Forewing (Fig. 117) oblong-oval with a somewhat truncate apex; costal break and rudimentary ptero- 
stigma usually absent; fine spinules present throughout all cells; vein Rs strongly curved towards the 
margin; vein M evenly curved; vein M 1+2 reaching to wing apex; cell cw la shortish but tall; vein Cu la 
strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male 
proctiger (Fig. 188) simple; paramere (Fig. 189) with inner tooth somewhat set back from apex and 
without thickened setae on basal posterior margin; aedeagus (Fig. 190) with strongly recurved apex. 
Female terminalia as in Fig. 290. 

Measurements. See Table 3. 

Material examined 

10 Cf, 11 $ , Switzerland and Italy (IDH, BMNH, DB). 

Host plant. Genista radiata (L.) Scop. 

Published distribution Austria (Low, 1888a; Prohaska, 1929; Haupt, 1935), Italy (Tamanini, 1977), 
Switzerland (Cerutti, 1939a,b; Schaefer, \949a,b; Burckhardt, 1983), Yugoslavia (Reuter, 1875). 

Comments. L. vittipennella is distinguished from other members of the genus by the extensive brown band 
extending along the fore margin of the wing. 



Livilla klapperiehisp. n. 

(Figs 155, 156, 230-232, 284) 

Description. Coloration. Genal cones, vertex and dorsum of thorax brick red with white markings; 
underside of head and thorax orange-yellow; abdomen brownish yellow; genitalia yellow; forewing 
membrane clear with reddish brown pattern; veins yellow to brown; antennal segments 1-8 dirty yellow, 
segments 4-8 apically infuscate; segments 9-10 dark brown; legs orange-yellow. 

Structure. Head (Fig. 156) with genal cones somewhat divergent, about as long as vertex, with narrowly 
rounded apices. Forewing (Fig. 155) oblong-oval, broadest at about two-thirds the length; costal break and 
rudimentary pterostigma present; fine spinules present throughout all cells; cell m x small; vein M 1+2 
reaching to wing apex; cell cw la relatively long, with vein Cw la relatively weakly curved; vein M evenly 
curved; vein Rs moderately strongly curved to wing margin. Metatibia with 4 thick black apical spurs; basal 
metatarsus with 2 black spurs. Male proctiger (Fig. 230) simple; paramere (Fig. 231) moderately stout, 
without basal posterior thickened setae; aedeagus (Fig. 232) with large bulbous apex. Female terminalia 
(Fig. 284) shorter and more wedge-shaped than in most other species. 

Measurements. See Table 3. 

Holotype <$ (on two slides), Afghanistan: Kamdesch, Bashguital, Nuristan, 2200 m, 17.vii.1952 
(Klapperich) (BMNH). 

Paratypes. 1 cf , 1 $ and one specimen lacking genitalia, same data as holotype. One female is on the 
same slide as the holotype, with the wing on a separate slide labelled paratype. 

Host plant. Unknown. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 29 

Comments. The name klapperichi is a manuscript name used by Vondracek. This species, which differs 
from other representatives of the genus in having 2 basal metatarsal spines and only 4 spines at the apex of 
the tibia. It is somewhat atypical and represents the easternmost record for the genus Livilla. 



The rad/ala-group 

Members of this group can be recognised by the strongly upturned apex of vein Rs and the vein A/, + 2 which 
reaches the wing margin well above the wing apex. 



Livilla radiata (Forster) comb. n. 
(Figs 98, 125,126,200-202) 

Arytaina radiata Forster, 1848: 70. Holotype $, Austria [not examined]. 

Psylla lactea Costa, 1863: 47; Low, 1877: 125; 1883: 241; Puton, 1876: 284. Type [sex not stated], Italy [not 
examined]. [Synonymised by Low, 1877: 125.] 

Psyllodes cytisi Becker, 1867: 113; Low, 1879: 595; 1883: 236. Type [incomplete data], U.S.S.R. [not 
examined]. [Synonymized by Low, 1879: 595.] 

Psylla radiata (Forster) Low, 1877: 125. 

Alloeoneura radiata (Forster) Low, 1879: 595; 1882a: 168; 1883: 248; 1884: 150; 1888a: 20; Horvath, 1886 
315; 1918: 59; Duda, 1892: 38; Aulmann, 1913: 35; Oshanin, 1912: 128; Sulc, 1905a: 4; Haupt, 1935: 241 
Prohaska, 1928a: 5; Klimaszewski, 1964: 65; 1965: 203; 1967: 13; 1969a: 46; 1973: 189; 1975: 132 
Loginova, 1964: 464; 1966: 135; 1976a: 599; Lauterer, 1963: 148; 1977: 98; Vondracek, 19516: 127; 1957 
189; Wagner & Franz, 1961: 162; Dobreanu & Manolache, 1962: 137; Andrianova & Klimaszewski, 
1983: 38; Glowacka & Harisanov, 1983: 64. 

Allaeoneura radiata (Forster); Puton, 1886: 93; Hueber, 1904: 276. Oshanin, 1907: 368. [Mis-spelling.] 

Aleuroneura radiata (Forster); Klimaszewski, 1961: 79. [Mis-spelling.] 

Description. Coloration. Mature specimens with head and thorax yellowish brown. Abdomen brown with 
yellow intersegmental membranes; genitalia yellow-brown; forewing membrane translucent white, veins 
yellow, markings chocolate brown; antennae yellow, apices of segments 3-8 brown, segments 9 and 10 
completely brown; legs yellowish brown; apices of tibiae and tarsi dark brown. 

Structure. Head (Fig. 126) with genal cones long and slender, much longer than the vertex. Forewing 
(Fig. 125) oblong-oval, very short and broad; costal break and pterostigma usually absent; fine spinules 
present throughout all cells; vein Rs short, strongly curved to wing margin; vein M short, slightly sinuous; 
vein M 1+2 meeting wing margin well above the apex; cell cw la short and relatively high, vein Cw la strongly 
arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 
200) simple; paramere (Fig. 201) without thickened setae along basal posterior margin; aedeagus (Fig. 202) 
with slightly reniform apex. Female terminalia as in Fig. 98. 

Measurements. See Table 3. 

Material examined 
2 cf, 3 $ , Austria, West Germany and Czechoslovakia (BMNH, DB). 

Host plants. Chamaecytisus ratisbonensis (Schaeffer) Rothm., Chamaecytisus austriacus (L.) Link, 
Chamaecytisus borysthenicus (Gruner) A. Klaskova, Lemboptropis nigricans (L.) Griseb. 

Published distribution. Austria (Forster, 1848; Low, 1863, 1888a; Prohaska, 1928a; Haupt, 1935; 
Wagner & Franz, 1961), Bulgaria (Klimaszewski, 1965; Glowacka & Harisanov, 1983), Czechoslovakia 
(Low, 1888a; Duda, 1892; Sulc, 1905a; Haupt, 1935; Vondracek, 1957; Lauterer, 1977), Hungary (Low, 
1888a; Horvath, 1886, 1918), Italy (Aulmann, 1913), Poland (Klimaszewski, 1961, 1964, 1967, 1969, 
1975), Rumania (Dobreanu & Manolache, 1962), U.S.S.R., southern European part (Loginova, 1964, 
1966; Andrianova & Klimaszewski, 1983), Yugoslavia (Low, 1888a; Aulmann, 1913), 'S. Germany' 
(Haupt, 1935). 

Comments. This species has been recognised as the type-species of the monotypic genus Alloeoneura Low, 
1879 for over 100 years. There is, however, little reason why it should be regarded as a distinct genus. L. 
radiata is distinguished by the short broad wing, with the characteristic pattern, and the very long genal 
cones. 



30 I. D. HODKINSON & D. HOLLIS 

Livilla retamae (Puton) comb. n. 
(Figs 119, 120, 191-193, 291) 

Psylla retamae Puton, 1878: 134; Bolivar & Chicote, 1879: 184. Syntypes 6 cf, 12 $, 1 ?sex, Spain (MM, 

MNHN) [examined] . 
Floria retamae (Puton) Low, 1883: 248; Puton, 1886: 93; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 

34; Ramirez Gomez, 1956: 85; Loginova, 1976a: 599; Klimaszewski, 1973: 194; Samy, 1973: 452; 

Halperinefa/.,1982:34. 
Floria (Brinckitia) retamae (Puton); Loginova, 1971: 629; 1972a: 27. 
AUoeoneura retamae (Puton) Vondracek, 1951ft: 127. 

Description. Coloration. Immature specimens greenish throughout, developing pale orange markings on 
dorsum of thorax. Mature specimens with genae, vertex and dorsum of thorax dirty yellow, with orange 
markings on thorax and vertex; ventral parts of head and thorax pale orange-yellow; dorsal abdominal 
sclerites brown, intersegmental membranes yellow; underside of abdomen greenish to brown; genitalia 
dirty yellow to brown; forewing membrane clear, veins clear to pale yellow; pattern brown; antennae dirty 
yellow, segments 3-6 darkened apically, segments 6-10 dark brown; legs dirty yellow. 

Structure. Head (Fig. 120) with genal cones slender, longer than the vertex, with narrowly rounded 
apices. Forewing (Fig. 119) elongate oblong-oval; costal break and rudimentary pterostigma present; 
spinules absent or thinly scattered in areas of brown patterning; vein Rs curved towards wing margin; vein 
M sinuous; vein M 1+2 meeting wing margin well before apex; cell cw la moderately large, vein Cw la strongly 
arched. Metatibia with 5 thick black spurs, basal metatarsus without black spurs. Male proctiger (Fig. 191) 
simple; paramere (Fig. 192) slightly expanded subapically, inner tooth partially hidden in side view; 
aedeagus (Fig. 193) with slightly hooked apex. Female terminalia as in Fig. 291. 

Measurements. See Table 3. 

Material examined 

16 Cf, 32$, Israel, Egypt, Algeria, Morocco, Portugal and Spain (IDH, ZI, ZMU, BMNH, MM, 
MNHN, DB, PL). 

Host plants. Retama sphaerocarpa (L.) Boiss, R. monosperma (L.) Boiss and R. raetam (Forskal) Webb 
& Berth. 

Published distribution. Algeria (Aulmann, 1913), Egypt (Samy, 1973), Israel (Halperin et al, 1982), 
Morocco (Loginova, 1972a), Portugal (Aulmann, 1913), Spain (Puton, 1878; Bolivar & Chicote, 1879; 
Vondracek, 19516; Ramirez Gomez, 1956). 

Comments. A widely distributed species which occurs right across the southern part of the Mediterranean 
Basin. It is distinguished from other members of the genus by the characteristic pattern and somewhat 
oblong shape of the forewing. 

Livilla lautereri sp. n. 

(Figs 121, 122, 194-196, 292) 

Description. Coloration. Available specimens do not appear to have developed the full mature coloration. 
General coloration of head and thorax brownish yellow, abdomen slightly darker above; genitalia brown; 
forewing membrane clear with very pale yellow clouds and very distinct dark brown markings towards 
apex; veins yellowish basally becoming dark brown at wing apex; antennae brownish yellow, segments 
becoming infuscate towards apex; legs yellowish brown. 

Structure. Head (Fig. 122) with genal cones widely separate and longer than vertex. Forewing (Fig. 121) 
oblong-oval, almost parallel-sided, with broadly rounded apex; spinules confined to apical cells and around 
base of vein Cu 2 ; costal break and pterostigma absent; vein Rs strongly curved to wing margin; vein M 
evenly curved; vein M 1+2 reaching wing margin well above apex; cell cw la moderately large, vein Cu ia 
straight in apical half. Metatibia with 4 or 5 thick black apical spurs; basal metatarsus with 1 black spur. 
Male proctiger (Fig. 194) simple; paramere (Fig. 195) with highly serrated anterior margin and large apical 
tooth; aedeagus (Fig. 196) very thin with angular apex. Female terminalia as in Fig. 292. 

Measurements. See Table 3. 

Holotype cf, Algeria: Massiv de l'Aures, 6 km Arris, 1300 m, 6.vi.l980 (Remane) (BMNH). 
Paratype. 1 $ , same data as holotype. 

Host plant. Unknown. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 31 

Comments. The forewing shape is similar to L. retamae but the pattern is quite different and characteristic, 
separating L. lautereri from other members of the genus. 

Livilla syriaca (Low) comb. n. 
(Figs 123, 124, 197-199,286) 

Floriasyriaca Low, 1882/?: 262; 1883: 252; Puton, 1886: 93; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 
35; Loginova, 1976a: 599; Halperin et al. , 1982: 34. Holotype $, Syria [not located, missing from Low 
collection (NM)]. 

Floria syriac(e) ] a Low r ; Klimaszewski, 1973: 194. [Mis-spelling.] 

Description. Coloration. Mature specimens with dorsum of thorax shining chocolate brown, remainder of 
thorax brownish yellow, occasionally with a greenish tinge; vertex and genal cones brownish yellow, 
occasionally greenish, marked along posterior and eye margins in darker brown; abdomen with dorsal 
sclerites brown, intersegmental membranes yellowish or greenish; ventral sclerites brown, occasionally 
greenish; genitalia yellow-brown to brown; forewing membrane clear to pale yellow, veins yellow to dark 
brown where they are overlaid with dark brown maculations; antennae yellowish brown, apices of 
segments 3-8 and whole of segments 9 and 10 brown. 

Structure. Head (Fig. 124) with genal cones triangular, widely divergent, about equal in length to vertex. 
Forewing (Fig. 123) oblong-oval but becoming broader towards the somewhat truncate apex; costal break 
and rudimentary pterostigma present; vein Rs strongly curved towards margin; vein M strongly sinous; 
vein M 1+2 meeting wing margin well above apex; cell cw la relatively tall, vein Cw la strongly curved. 
Metatibia with 5 thick black spurs; basal metatarsus with 1 spur. Male proctiger (Fig. 197) simple; 
paramere (Fig. 198) long and slender without thickened setae on basal posterior margin; aedeagus (Fig. 
199) bulbous and slightly recurved at apex. Female terminalia as in Fig. 286. 

Measurements. See Table 3. 

Material examined 
2 cT, 13 $, Cyprus (BMNH, MM). 

Host plant. Genista fasselata Decne and Genista acanthoclada D.C. 

Published distribution. Syria (Low, 1882b) and possibly Israel (Halperin etal. , 1982). 

Comments. The types of this species are missing from the Low collection in the NM, Vienna. We have 
followed Loginova's interpretation of the species which is, in all probability, correct. This species is 
distinguished by the forewing, which is strongly narrowed toward the base, and by the widely divergent 
genal cones. 

The spartiisuga-group 

Members of this group are characterised by the forewing pattern consisting primarily of small round 
maculations. 

Livilla spartiisuga (Puton) comb. n. 

(Figs 137, 138, 218-220, 276) 

Psylla spartiisuga Puton, 1876: 283. Lectotype cf , Algeria (MNHN), here designated [examined]. 
Floria spartiisuga (Puton) Low, 1879: 593; 1883: 251; Puton, 1886: 92; Oshanin, 1907: 367; 1912: 128; 
Aulmann, 1913: 34; Loginova, 1971: 629; 1976a: 599. 

Description. Coloration. Mature specimens with dorsum of thorax orange-red with brownish markings; 
head including genal cones orange ; abdominal sclerites brown , intersegmental membranes reddish orange ; 
genitalia orange-brown to dark brown; forewing membrane clear to pale yellow, veins yellow, maculations 
pale brown; antennae yellow with apices of segments black; legs yellowish orange. 

Structure. Head (Fig. 138) with genal cones slightly longer than vertex, stout and broadly rounded. 
Forewing (Fig. 137) elongate oblong-oval; with costal break and rudimentary pterostigma present; 
spinules present throughout all cells; vein Rs moderately strongly curved towards margin; vein M evenly 
curved throughout; vein A/ 1+2 reaching to wing apex; cell cw la moderately long and flat. Metatibia with 5 
thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 218) simple; paramere 
(Fig. 219) long and slender; aedeagus (Fig. 220) with rounded bulbous apex. Female terminalia as in Fig. 
276. 



32 I. D. HODKINSON & D. HOLLIS 

Measurements. See Table 3. 

Material examined 

Lectotype cf , Algeria: Bone, with Puton type label (MNHN). 2 cf , 4 £ paralectotypes, same data as 
lectotype (MNHN). 

Host plant. Genista sp. 

Published distribution. Algeria (Puton, 1876). 

Comments. This species is close to siciliensis and known only from the type series. It can be separated from 
all other 'spotted-wing' species by the absence of maculations from a band stretching primarily across cell r s 
of the fore wing. 

Livilla siciliensis sp. n. 

(Figs 131, 132, 209-211, 273) 

Description. Coloration. Immature specimens pale greenish to yellow throughout. Some of the more 
mature specimens are more yellowish and are beginning to develop orange markings on the dorsum of the 
thorax and vertex and the abdominal sclerites are beginning to darken. Genitalia yellow to brown; 
fore wing membrane clear to pale yellowish, veins yellowish to pale brown; maculations brown; antennal 
segments 1-4 yellow, segments 3-4 infuscate at apex; segments 5-10 dark brown; legs dirty yellow. 

Structure. Head (Fig. 132) with genal cones rather stout, about as long as vertex, with broadly rounded 
apices. Forewing (Fig. 131) oblong-oval, almost parallel-sided; costal break and rudimentary pterostigma 
present; fine spinules present throughout all cells; vein R s curved towards margin; vein M long and very 
slightly sinuous; cell m 1+2 small. Vein M 1+2 reaching to wing apex; cell cw la relatively flat. Metatibia with 5 
thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 209) simple; paramere 
(Fig. 210) long and slender; aedeagus (Fig. 211) with slightly angular bulbous apex. Female terminalia as in 
Fig. 273. 

Measurements. See Table 3. 

Holotype cf , Italy: Sicily, Palermo c. 750 m, 28.V.1979 (D. & S. Sutton) (BMNH). 
Paratypes. 10 cf, 18 $, same data as holotype except that some were collected on 26. v. 1979 and 
5.vi.l979. 

Host plant. Genista sp. 

Comments. Morphologically this species is closest to L. maculipennis but it is much larger. 

Livilla cataloniensis (Hodkinson & White) comb. n. 

(Figs 127, 128, 203-205, 271) 

Floria cataloniensis Hodkinson & White, 1979a: 55. Holotype cf , France (BMNH) [examined]. 

Description. Coloration. Mature specimens with dorsal surface of head and thorax orange-yellow with 
paler longitudinal markings; genal cones and underside of thorax brownish yellow; abdominal sclerities 
brown, often darker above; intersegmental membranes yellow, occasionally with greenish tinge in younger 
specimens; genitalia yellow to yellow-brown; forewing membrane clear to very pale yellow throughout, 
maculae brown; antennal segments 1-2 yellow; segments 3-7 yellow with infuscate apex; segments 9-10 
dark brown; legs brownish yellow. 

Structure. Head (Fig. 128) with genal cones conical, slightly longer than the vertex, with narrowly 
rounded apices. Forewing (Fig. 127) oblong-oval with broadly rounded apex; pterostigma rudimentary; 
costal break occasionally present; fine spinules present throughout all cells; vein Rs weakly curved towards 
margin; vein M evenly curved; vein M 1+2 reaching to wing apex; cell cw la short and high; vein Cu la strongly 
arched. Metatibia with 5 thick black apical spurs at apex; basal metatarsus with 1 black spur. Male proctiger 
(Fig. 203) simple; paramere (Fig. 204) relatively stout and straight, without thick basal setae along 
posterior margin; aedeagus (Fig. 205) with bulbous apex. Female terminalia as in Fig. 271. 

Measurements. See Table 3. 

Material examined 

Cf holotype, France: Pyrenees-Or., Argeles, 16.V.1977 (White) (BMNH). 15 cf, 21 $ including 
paratypes, France and Spain (BMNH, IDH). There is also a lot of material from unnamed localities in the 
Heslop-Harrison collection (BMNH). 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 33 

Host plant. Ulex parviflorus Pourret. 

Published distribution. France and Spain (Hodkinson & White, 1979a). 

Comments. This species can be distinguished from other spotted-wing species by the relatively broader 
fore wing. 

Livilla hodkinsoni (Burckhardt) comb. n. 

(Figs 139, 140,221-223,277) 

Floria hodkinsoni Burckhardt, 1979: 391. Holotype Q\ Greece (ETH) [examined]. 

Description. Coloration. Immature specimens bright green throughout with orange markings on dorsum 
of head and thorax. Mature specimens with dorsal surface of head and thorax brown with yellow markings; 
underside of head and thorax with dark brown markings; abdominal sclerites dark brown, intersegmental 
membranes paler; genitalia yellow-brown to brown; forewing membrane clear to pale yellow with brown 
maculations; antennal segments 1-7 yellow, segments 4-7 infuscate at apex; segments 8-10 dark brown; 
legs brownish yellow. 

Structure. Head (Fig. 140) with genal cones longer than vertex, with somewhat angular apices. Forewing 
(Fig. 139) oblong-oval, with rudimentary pterostigma and costal break present; fine spinules present 
throughout all cells; vein R s moderately strongly curved towards margin; vein M slightly sinuous; vein M l+2 
reaching to just below the wing apex; cell cw la moderately long, of characteristic shape. Metatibia with 5 
thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 221) simple; paramere 
(Fig. 222) without thickened basal posterior setae; aedeagus (Fig. 223) with bulbous apex. Female 
terminalia as in Fig. 277. 

Measurements. See Table 3 

Material examined 

Cf holotype, Greece: Pelopones, Bilos, 16. v. 1979 (Burckhardt) (ETH). 13 d\ 15 $ including paratypes, 
Greece, including Crete (ETH, DB, BMNH, MM, PL). 

Host plant. Genista acanthoclada D.C., not a Cytisus species as stated in the original description 
(Burckhardt, pers. comm.). 

Published distribution. Greece (Burckhardt, 1979). 

Comments. L. hodkinsoni is closest to L. cataloniensis from which it is separated by the shape of cells m x 
and cw la of the forewing. 

Livilla maculipennis sp. n. 

(Figs 129, 130, 206-208, 272) 

Description. Coloration. Body colour of immature specimens bright green to yellow, usually with orange 
or orange-brown markings on dorsum of vertex and thorax. Mature specimens darker, general colour pale 
brown with darker brown markings; abdominal sclerites dark brown with yellow intersegmental mem- 
branes; forewing membrane clear, veins pale yellow, maculations brown; antennal segments brownish 
yellow basally becoming dark brown towards apex. 

Structure. Head (Fig. 130) with genal cones divergent, narrowly rounded at apex, slightly longer than 
vertex. Forewing (Fig. 129) elongate, oblong-oval, almost parallel-sided, pterostigma almost lost, costal 
break often present; fine spinules present throughout all cells; vein Rs moderately strongly curved towards 
wing margin; vein M slightly sinuous; vein M 1+2 reaching to wing apex; cell cu la very long and low. 
Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 206) 
simple; paramere (Fig. 207) curved posteriorly beyond middle, with numerous thickened setae on basal 
posterior margin; aedeagus (Fig. 208) with apex somewhat reniform. Female terminalia as in Fig. 272. 

Measurements. See Table 3. 

Holotype cf , Spain: Huelva, 26 km, S. Almonte, 13. vi. 1977 (Hollis) (BMNH). 

Paratypes. Spain: 11 cf, 11 $, same data as holotype. Algeria: 7 cf , 3 $, Theniet al Had, Zedernwald, 
1700 m, 31.vii.1980 {Remane) (PL); 10 cf, 6 ?, NE1 Gor, 900 m, 28.vii.1980 (Remane) (PL); 7 cf , 11 ?, 
Monte de Tlemcen, 18 km NW. Sebdou, 27.vii.1980 (Remane) (PL). Morocco: 1 cf, N Taferaet, 
25.vii.1980 (Remane) (PL). 

Host plant. Genista sp. 



34 I. D. HODKINSON & D. HOLLIS 

Comments. L. maculipennis is separated from the other spotted-wing species by its small size and the 
elongate cell cw la of the forewing. 

Livilla pyrenaea (Mink) comb. n. 

(Figs 133, 134, 212-214, 274) 

Psylla pyrenaea Mink, 1859: 430. Syntypes, France [not located]. 

Floria pyrenaea (Mink) Low, 1879: 592; 1883: 247; Chicote, 1880: 202; Puton, 1871: 438; 1886: 92; 

Oshanin, 1907: 367; 1912: 128; Aulmann, 1913: 34; Vondracek, 19516: 128; Loginova, 1976a: 599; 

Hodkinson & White, 1979a: 57. 
Floria (Floriella) pyrenaea (Mink); Ramirez Gomez, 1956: 87. 
Floria pyrenea (Mink); Klimaszewski, 1973: 194. [Mis-spelling.] 

Description. Coloration. Mature specimens with dorsal surface of head and thorax dark orange or 
orange-brown with paler longitudinal markings; genal cones and underside of thorax brownish yellow; 
abdominal sclerites dark brown with yellow intersegmental membranes, often with a greenish tinge in less 
mature specimens; genitalia yellow to brown; forewing membrane clear; maculae brown, absent from area 
adjacent to wing margin, making leading edge of wing appear white from a distance; antennal segments 
1-2 yellow; segments 3-7 yellow with infuscate apex; segments 9-10 brown; legs brownish yellow. 

Structure. Head (Fig. 134) with genal cones somewhat triangular, divergent, longer than vertex. 
Forewing (Fig. 133) oblong-oval with a somewhat truncate apex; costal break and pterostigma usually 
absent; fine spinules present throughout all cells; vein Rs curved towards wing margin; vein M evenly 
curved; vein M 1+2 reaching to wing apex; cell cw la moderately long. Metatibia with 5 thick black apical 
spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 212) simple; paramere (Fig. 213) relatively 
short, parallel-sided; aedeagus (Fig. 214) robust, with slightly reniform apex. Female terminalia as in Fig. 
274. 

Measurements. See Table 3. 

Material examined 
20 cf , 19 ? , France and Spain (BMNH, IDH, MM). 

Host plants. Calicotome spinosa (L.) Link, and Genista pilosa L. 

Published distribution. Spain (Chicote, 1880; Ramirez Gomez, 1956; Hodkinson & White, 1979a), 
France (Mink, 1859; Puton, 1871; Vondracek, 19516, Hodkinson & White, 1979a). 

Comments. A characteristic species with maculations absent from the leading margin of the forewing. It 
also differs from other spotted-wing species by having a slightly obliquely truncated wing apex. 

Livilla bimaculata sp. n. 

(Figs 149, 150, 254-256, 280) 

Description. Coloration. Mature specimens with dorsal surface of head and thorax pale reddish brown 
with paler markings; genal cones and ventral parts of thorax brownish yellow; abdominal sclerites dark 
brown; intersegmental membranes yellowish; genitalia brown; forewing membrane two-tone, clear and 
yellow-brown; maculae generally mid brown, those around the apex of vein Rs darker, almost black; 
antennae with segments 1-4 primarily yellow, segments 6-10 primarily dark brown; legs brownish yellow. 

Structure. Head (Fig. 150) with genal cones somewhat divergent, as long as vertex. Forewing (Fig. 149) 
elongate oval, broadest in basal half; fine spinules present throughout all cells; vein Rs strongly curved 
towards wing margin; vein M sinuous; vein M i+2 reaching to wing apex; pterostigma at most rudimentary; 
costal break occasionally present. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black 
spur. Male proctiger (Fig. 254) simple; paramere (Fig. 255) slender, with numerous stout spines on basal 
posterior margin; aedeagus as in Fig. 256. Female terminalia as in Fig. 280. 

Measurements. See Table 3. 

Holotype cf, France: Corsica, Gorge du Ristomica nr Corte, 12. vi. 1976, on Genista sp. (J.M.P.) 
(BMNH). 

Paratypes. 6 cf, 10 J, same data as holotype. Italy: 4 cf, 4 J, Sardinia, Genova Borzonasea, Passo, 
Boco (Conci) 16.vi.1984 (IDH). 

Host plant. Genista salzmanii DC. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 35 

Comments. This species is separated from other 'spotted-wing' Livilla species by the presence of a much 
darker patch of maculations around vein Rs. 

Livilla blandula (Horvath) comb. n. 
(Figs 144-146, 236-238, 278) 

Floria blandula Horvath, 1905: 277; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 34; Ramirez Gomez, 
1956: 84; Klimaszewski, 1973: 193; Loginova, 1976a: 599. LECTOTYPE cf, Spain (TM) here 
designated [examined]. 

[Floria maura Vondracek sensu Loginova, 1972a: 27. Misidentification.] 

Description. Coloration. Body colour of immature specimens brownish yellow. Mature specimens with 
dorsum of thorax and vertex chocolate brown with white markings; genal cones and underside of thorax 
brownish yellow; dorsum of abdomen brown, remainder pale green to yellow; genitalia greenish yellow to 
brown; fore wing membrane two-tone, clear and yellow-brown; maculae mid to dark brown; antennae with 
segments 1-5 primarily yellow, segments 6-10 primarily brown. 

Structure. Head (Fig. 146) with genal cones slightly longer than vertex, slightly divergent, with apices 
narrowly rounded. Forewing (Figs 144, 145) oblong-oval; costal break and rudimentary pterostigma often 
present; fine spinules present throughout all cells; vein Rs weakly curved towards wing margin; vein M 
slightly sinuous; vein M 1+2 reaching to wing apex. Metatibia with 5 thick black apical spurs; basal 
metatarsus with 1 black spur. Male proctiger (Fig. 236) simple; paramere (Fig. 237) with numerous thick 
setae on basal posterior margin; aedeagus (Fig. 238) with slightly hooked apex. Female terminalia as in Fig. 
278. 

Measurements. See Table 3. 

Material examined 
Lectotype O" , Spain: 'Hispania. Ciudad Real, det. Horvath' (TM). 
1 0\ 4 $ (paralectotypes), same data as lectotype; 19 cf , 27 $ , Portugal, Morocco (BMNH, ZMU, ZI). 

Host plant. Genista hirsuta Vahl. 

Published distribution. Spain (Horvath, 1905), Morocco (Loginova, 1972a). 

Comments. This is the species illustrated as maura by Loginova (1972a). The forewing of maura is 
relatively broader and with only scattered indistinct maculations. This species shows considerable variation 
in the wing pattern and the two extremes are illustrated for comparison. Livilla blandula is closest to L. 
lusitanica and L. pseudoretamae . It can be distinguished from the former by the narrower forewing and 
from the latter by the lack of a distinct dark brown forewing pattern. 

Livilla pseudoretamae sp. n. 

(Figs 157, 158, 233-235, 285) 

Description. Coloration. Vertex, genal cones and dorsum of thorax pale greyish brown, occasionally with 
a greenish tinge, thorax with chocolate brown markings; abdominal sclerites ranging from green to brown, 
with pale intersegmental membranes; genitalia dirty yellow to brown; forewing membrane clear, veins 
yellow to very pale brown; wing pattern light chocolate-brown; antennae dirty yellow basally, segments 
5-10 dark brown; legs brownish yellow throughout. The specimens to hand have probably not developed 
their full coloration. 

Structure. Head (Fig. 158) with genal cones slightly longer than vertex, with evenly rounded apices. 
Forewing (Fig. 157) oblong-oval; with costal break and rudimentary pterostigma present; fine spinules 
present throughout all the cells; vein Rs moderately strongly curved towards margin; vein M moderately 
sinuous; vein M 1+2 short but reaching to wing apex; cell cw la short and high, vein Ca la strongly arched. 
Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 233) 
simple; paramere (Fig. 234) straight, with thickened setae present on basal posterior margin; aedeagus 
(Fig. 235) with a slightly hooked apex. Female terminalia as in Fig. 285. 

Measurements. See Table 3. 

Holotype cf, Algeria: Gran Kabylie, Djurdjura, 3.viii.l980 (Remane) (BMNH). 
Paratypes. 2 cf , 5 $>, same data as holotype; 3 cf, 9 $, Kabylie, Col de Talmetz {Remane). (BMNH, 
PL.) 

Host plant. Unknown. 



36 I. D. HODKINSON & D. HOLLIS 

Comments. This species is very close to L. blandula but the wing pattern, which lacks the characteristic 
spots, is more distinct and superficially resembles that of Livilla retamae. 

Li villa lusitanica sp. n. 

(Figs 147, 148, 251-253, 279) 

Description. Coloration. Mature specimens with dorsum of head and thorax brown with whitish markings; 
genal cones and underside of thorax brownish yellow; abdominal sclerites dark brown, intersegmental 
membranes yellowish brown; genitalia brown; fore wing membrane two-tone, clear and yellow-brown; 
maculae mid to dark brown; antennae with segments 1-5 primarily yellow, segments 6-10 primarily dark 
brown; legs brownish yellow. 

Structure. Head (Fig. 148) with genal cones stout, about as long as vertex. Fore wing (Fig. 147) broadly 
oblong-oval, with costal break and rudimentary pterostigma present; fine spinules present throughout all 
cells; vein Rs moderately strongly curved towards wing margin; vein M slightly sinuous; vein M 1+2 reaching 
wing apex; cell cu la strongly curved. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black 
spur. Male proctiger (Fig. 251) simple; paramere (Fig. 252) stout, with dense thickened setae on the basal 
posterior margin; aedeagus (Fig. 253) with apex somewhat reniform. Female terminalia as in Fig. 279. 

Measurements. See Table 3. 

Holotype C\ Portugal: 29 km E. Coimbra, 21.vi.1977 (Hollis) (BMNH). 

Paratypes. 7 cf, 10 $, same data as holotype; 3 cf, 3 $, Porto, 2-12. viii. 1962 (/. Abraham & L. 
Horascek) (BMNH). 

Host plant. Genista triacanthos Brot. 

Comments. This species resembles blandula in several respects but the fore wing is consistently shorter and 
broader and the genal cones slightly shorter. 

Livilla magna sp. n. 

(Figs 135, 136, 215-217, 275) 

Description. Coloration. Immature specimens pale green to yellow throughout. Mature specimens with 
vertex and dorsum of thorax brownish yellow with orange or chocolate brown markings; underside of 
thorax often with most of sclerites dark brown to black; genal cones brownish yellow; sclerites of abdomen 
dark brown, occasionally paler along ventral parts; intersegmental membranes brownish yellow; forewing 
two-tone, clear to pale yellow contrasted with amber; maculations medium brown; antennal segments 1-4 
yellow, segments 3-4 infuscate at apex; segments 5-10 dark brown; legs brownish yellow, femora 
occasionally darkened above. 

Structure. Head (Fig. 136) with divergent triangular genal cones as long as vertex. Forewing (Fig. 135) 
elongate oblong oval; rudimentary pterostigma and costal break present; vein Rs moderately curved to 
margin; fine spinules present in all cells; cells cu XaL and m 1+2 of average size for genus; vein M 1+2 reaching to 
wing apex. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male proctiger 
(Fig. 215) simple; paramere (Fig. 216) long and thin with a kink in the middle; aedeagus (Fig. 217) long 
with an angular bulbous apex. Female terminalia as in Fig. 275. 

Measurements. See Table 3. 

Holotype cf , Italy: Sicily, Catania, l.vi.1979 (D. & S. Sutton) (BMNH). 

Paratypes. 5 cf , 9 $ , same data as holotype; 2 cf, 8 $ , same data as holotype except 4.vi.l979; 1 cf , 7 $, 

Sicily, Etna, 1100 m, 16.vii.1977 (D'llrso) (PL); 1 cT, 3 $ , Sicily, Etna, 1100 m, 19.ix.1977 (Asche) (PL). 

Host plant. Genista aetnensis (Biv.) DG. 

Comments. This species is much larger than the other spotted-wing species. Morphologically it is closest to 
L. siciliensis but it differs in the shape of the forewing and details of the male genitalia. 

Livilla poggii (Conci & Tamanini) comb. n. 

(Figs 141-143, 224-226, 281) 

Floria poggii Conci & Tamanini, 1984a: 43. Holotype d\ Italy: Sardinia (MCSN) [not examined]. 

Description. Coloration. General coloration dirty yellow to greyish yellow throughout, occasionally with 
brown markings on the dorsum of the thorax; genitalia yellow to dark brown; forewing membrane clear to 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 37 

yellowish but covered in brown maculations which are often coalescent; in the female the maculations are 
spread throughout the wing whereas in the male there is a transverse maculation-free band across the 
middle; also in the latter the apical maculations appear darker than the basal ones; antennal segments 1-8 
yellow, segments 3-8 apically infuscate; segments 9-10 dark brown; legs orange-yellow. 

Structure. Head (Fig. 143) with genal cones weakly divergent, with narrowly rounded apices, about as 
long as vertex. Forewing (Figs 141 , 142) elongate oval, more so in the female; costal break and pterostigma 
usually absent; fine spinules present in all cells; vein R s strongly curved towards wing margin; vein M 
strongly sinuous; vein M 1+2 meeting margin just above wing apex; cell cu Vd short and tall; vein Cw la 
strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black spur. Male 
proctiger (Fig. 224) simple; paramere (Fig. 225) relatively short and stout; aedeagus (Fig. 226) with an 
irregularly angled apex. Female terminalia as in Fig. 281. 

Measurements. See Table 3. 

Material examined 
2 cf , 3 $ (paratypes), Italy (IDH). 

Host plant. Genista Corsica (Loisel.) DC in Lam. & DC. 

Published distribution. Italy, Sardinia (Conci & Tamanini, 1984a). 

Comments: This is an interesting species which is strongly sexually dimorphic in the wing-pattern, an 
unusual occurrence in the psyllids. The host plant is endemic to Sardinia and Corsica. 



Residual assemblage 

Livilla spectabilis (Flor) comb. n. 

(Figs 161, 162, 242-244, 268) 

Psylla spectabilis Flor, 1861: 362. Syntypes d\ $ France (not located). 

Floria spectabilis (Flor) Low, 1879: 594; 1883: 251: 1888a: 20; Chicote, 1880: 202; Ferrari, 1888: 76; Puton, 

1886: 93; Oshanin, 1907: 368; 1912: 128; Aulmann, 1913: 34; Horvath, 19186: 331; Haupt, 1935: 242; 

Schaefer, 1949a: 56, 19496: 31; Vondracek, 19516: 128; Klimaszewski, 19686: 781; 1973: 194; Novak & 

Wagner, 1962: 43; Loginova, 1976a: 599; Burckhardt, 1983: 54. 

Description. Coloration. Immature specimens pale green to greenish yellow throughout. Mature speci- 
mens with genal cones, vertex and dorsum of thorax dirty orange-yellow with darker orange and paler 
streaks; abdominal sclerites dark brown, intersegmental membranes orange-yellow; genitalia orange- 
yellow to brown; forewing membrane clear basally becoming orange-brown apically, with a dark spot 
around vein Cw la ; antennae dirty yellow, except segments 3 and 4 apically darkened, segments 5 to 10 
brown; legs dirty yellow. 

Structure. Head (Fig. 162) with massive genal cones which are much longer than the vertex. Forewing 
(Fig. 161) elongate oval, of characteristic shape, with a strongly curved costal margin throughout; costal 
break and rudimentary pterostigma normally absent; fine spinules present throughout all cells; vein Rs 
weakly curved towards margin; vein M evenly curved; vein M 1+2 reaching to wing apex; cell cw la relatively 
small; vein Cw la strongly arched. Metatibia with 5 thick black apical spurs; basal metatarsus with 1 black 
spur. Male proctiger (Fig. 242) simple; paramere (Fig. 243) straight and slender; aedeagus (Fig. 244) with 
recurved apex. Female terminalia as in Fig. 268. 

Measurements. See Table 3. 

Material examined 

10 d\ 9 $, Italy, Greece including Crete, Yugoslavia, France (MM, BMNH, IDH, DB, PL). There is 1 
CT, 1 2 in the ZI, Leningrad labelled Uppsala, Sweden, F. Ossiannilsson. Dr Ossiannilsson informs us that 
this is a mistake; the species does not occur in Sweden. 

Host plant. Spartium junceum L. 

Published distribution. France (Puton, 1871), Italy (Ferrari, 1888), Portugal (Aulmann, 1913), Spain 
(Chicote, 1880), Switzerland (Schaefer, 1949a,6; Burckhardt, 1983). 

Comments. This species has no known close relative within the genus. It is separated from other members 
by the narrow forewing of characteristic shape and the very long genal cones. 



38 I. D. HODKINSON & D. HOLLIS 

Livilla variegata (Low) comb. n. 
(Figs 163, 164, 245-247, 269) 

Floria variegata Low, 18826: 261; 1883: 253; 1888a: 20; Ferrari, 1888: 76; Puton, 1886: 92; Oshanin, 1907 
367, 1912: 128; Aulmann, 1913: 35; Klimaszewski, 1973: 194; Loginova, 1976a: 599; Hollis, 1978: 149 
Hodkinson & Hollis, 1980: 171; Hodkinson & White, 1979a: 63; White & Hodkinson, 1982: 25 
Burckhardt, 1983: 54. Syntypes, cf $, Yugoslavia (MM) [examined]. 

Floria alpina Cerutti, 1939a: 447; 19396: 583; Schaefer, 1949a: 55; 19496: 31; Klimaszewski, 1973: 193; 
Loginova, 1976a: 599; Tamanini, 1977: 111. Lectotype cf, Switzerland (MZL) [not examined]. 
[Synonymised by Burkchardt, 1983: 54.] 

Description. Coloration. Immature specimens pale green to greenish yellow with orange markings on 
dorsum of thorax. Mature specimens with dorsum of head and thorax orange-yellow with paler longitudi- 
nal markings; genal cones orange-yellow; abdominal sclerites dark brown, intersegmental membranes 
yellow; genitalia orange-brown; fore wing membrane clear to pale yellow basally, apical suffusion grey to 
pale yellowish brown, veins pale yellow to very pale brown; antennae dirty yellow, apices of segments 3 to 6 
and whole of segments 7 to 10 dark brown; legs dirty yellow. 

Structure. Head (Fig. 164) with genal cones slender, slightly longer than vertex, with narrowly rounded 
apices. Forewing (Fig. 163) oblong-oval; costal break and rudimentary pterostigma present; veins very 
delicate; dense fine spinules present throughout all cells; vein Rs weakly curved to margin; vein M evenly 
curved; vein M 1+2 reaching to wing apex; cell cw la strongly arched. Metatibia with 5 thick black apical 
spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 245) simple; paramere (Fig. 246) slender, 
with thickened setae present on basal posterior margin; aedeagus (Fig. 247) with bulbous apex. Female 
terminalia as in Fig. 269. 

Measurements. See Table 3. 

Material examined 

27 cf, 32 $, Yugoslavia, Rumania, Italy, Switzerland, France, Spain, England (MM, IDH, BMNH, 
DB). 

Host plants. Laburnum anagyroides Medicus and Laburnum alpinum (Miller) Berchtold & J. Presl. 

Published distribution. France (Hodkinson & White, 1979a), Great Britain (Hollis, 1978; Hodkinson & 
Hollis, 1980; White & Hodkinson, 1982), Italy (Ferrari, 1888; Tamanini, 1977), Switzerland (Cerruti, 
1939a,6; Schaefer, 1949a,6; Burckhardt, 1983), Yugoslavia (Low, 18826; 1888a). 

Comments. Floria alpina was, for a long time, recorded as a separate alpine species but it has now been 
shown to be synonymous with L. variegata. L. variegata can be separated from other members of the genus 
by the broad forewing which lacks a distinct pattern and by the veins which are unusually fine and lightly 
coloured. 

Livilla genistae Ramirez Gomez 

(Figs 153, 154, 227-229, 283) 

Livilla genistae Ramirez Gomez, 1956: 98; Klimaszewski, 1973: 195; Loginova, 1976a: 600. NEOTYPE cf, 
Spain (BMNH) here designated. 

Description. Coloration. Mature specimens with head and thorax yellowish brown with slightly darker 
reddish brown markings; abdominal sclerites occasionally darker than thorax, terminalia yellow-brown to 
brown; forewing membrane clear, without pattern; veins yellowish brown; antennal segments 1-2 yellow, 
segments 3-4 yellow and infuscate at apex; segments 5-10 dark brown; legs brownish yellow. 

Structure. Head (Fig. 154) with genal cones almost as long as vertex, somewhat divergent with broadly 
rounded apices. Forewing (Fig. 153) elongate oval without pterostigma or costal break; spinules present 
throughout all cells; vein Rs strongly curved towards wing margin; vein M evenly curved; vein M 1+2 
reaching to just below wing apex; cell cw la short and tall; vein Cw la strongly curved. Metatibia with 5 apical 
thick black spurs; basal metatarsus with 1 black spur. Male proctiger (Fig. 227) simple; paramere (Fig. 228) 
sinuous, with thick setae at base of posterior margin; aedeagus (Fig. 229) with weakly angled apex. Female 
terminalia (Fig. 283) with very short circumanal pore ring. 

Measurements. See Table 3. 

Material examined 
Neotype cf , Spain: Huesca, Barluenga, l.vi.1976, on ? Genista (Hollis) (BMNH); 8 $ , Spain (BMNH). 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 39 

Host plant. This was omitted from the original description: the name genistae suggests that it feeds on a 
'Genista species. The neotype designated above was collected from a plant thought to be a Genista species 
while 4 of the additional females were collected from Cytisus purgans. 

Published distribution. Spain (Ramirez Gomez, 1956). 

Comments. The original type-material of this species has been destroyed. The above material collected in 
Spain corresponds closely with the original description. This species is somewhat intermediate between the 
short coriaceous wing Livilla species (ulicis-group) and the longer wing forms with spotted or longitudi- 
nally striped wing patterns (spartiisuga- and horvathi-groups) . It is recognised by the oval wing and the 
absence of any wing patterning. 

Livilla adusta (Low) comb. n. 
(Figs 159, 160,239-241,267) 

Floria adusta Low, 18826: 260; 1883: 230; Puton, 1886: 92; 1889: 113; Oshanin, 1907: 367; 1912: 128; 
Aulmann, 1913: 34; Klimaszewski, 1973: 193; Loginova, 1976a: 599. ? Holotype $, Spain [missing from 
Low collection (NM)]. 

Description. Coloration. Immature specimens greenish yellow throughout, developing pale orange 
markings on dorsum of thorax. Mature specimens with genal cones, vertex and dorsum of thorax brownish 
orange with indistinct brown or paler markings; abdominal sclerites dark brown to black, intersegmental 
membranes orange-brown. Forewing two-tone, clear and orange-brown; veins yellow to yellow-brown; 
antennae dirty yellow, segments 3-4 darkened apically, segments 5-10 dark brown; legs dirty yellow. 

Structure. Head (Fig. 160) with genal cones as long as vertex, slightly divergent with narrowly rounded 
apices; forewing (Fig. 159) elongate oblong-oval, with evenly rounded apex; costal break and pterostigma 
rudimentary; fine spinules present throughout all cells; veins Rs upturned to wing margin; vein M 1+2 
reaching to wing apex. Metatibia with 5 thick black spurs; basal metatarsus with 1 black spur. Male 
proctiger (Fig. 239) simple but broadest at base; paramere (Fig. 240) long, thin, parallel-sided, with large 
inner apical tooth, bearing a few stout setae at base of posterior margin; aedeagus as in Fig. 241. Female 
terminalia (Fig. 267) with a slightly upturned apex to the proctiger. 

Measurements. See Table 3. 

Material examined 
9 cf , 12 $ , Algeria and Spain (BMNH, MNHN, MM). 

Host plant. Unknown. Some specimens in BMNH were collected from Cytisus scoparius ? 

Published distribution. Spain (Low, 18826; Aulmann, 1913). 

Comments. This species is not very close to any other member of the genus. It is easily recognised by the 
distribution of darker pigment along the forewing veins which gives a furrowed appearance. 

Livilla maura (Vondracek) comb. n. 

(Figs 151, 152, 282) 

Floria maura Vondracek, 1951a: 119; 19516: 128; Klimaszewski, 1973: 193; Loginova, 1976a: 599. 
Syntypes $, Morocco (MM) [examined]. 

Description. Coloration. Mature specimens with vertex and dorsum of thorax deep brownish orange with 
darker brown markings; genal cones brown; abdomen dark brown to black, intersegmental membranes 
brownish orange; genitalia brownish yellow to dark brown; forewing membrane, pale brownish orange 
throughout, veins concolorous with membrane; apical pattern very diffuse, pale brown; antennae with 
segments 1-4 dirty yellow, 3-4 apically brown, the remaining segments brown; legs with femora pale 
brown, the remainder orange-yellow. 

Structure. Head (Fig. 152) with genal cones about as long as vertex, relatively slender, with narrowly 
rounded apices. Forewing (Fig. 151) oblong-oval, relatively broad; costal break and rudimentary ptero- 
stigma present; fine spinules present throughout all cells; vein M evenly curved; vein M l+2 reaching to wing 
apex; cell cu x relatively short with vein Cw la strongly arched. Metatibia with 5 thick black apical spurs; basal 
metatarsus with 1 black spur. Female terminalia as in Fig. 282. Male unknown. 

Measurements. See Table 6. 



40 



I. D. HODKINSON & D. HOLLIS 



Material examined 

2 $ (syntypes), Morocco (MM). In addition, there are 2 heads, 5 wings, 2 female genitalia and other 
oddments on slides labelled 63-69 Floria maura type series (MM). 

Host plant. Unknown. 

Published distribution. Morocco (Vondracek, 1951a, b). 

Comments. This species was described from several females. The male is unknown and this makes it 
difficult to characterize the species. It appears to differ from other members of the genus in possessing a 
relatively broad fore wing of uniform brownish orange coloration. 



Table 3 

mm. 



Measurements of Livilla species. Males and females are given separately and all values are in 



Head Antennal Forewing 
width length length 



Female Male Male Male 

proctiger proctiger paramere aedeagus 

length length length length 



adusta 


m 


0-63-0-74 




f 


0-67-0-74 


bimaculata 


m 


0-70-0-73 




f 


0-72-0-76 


bivittata 


m 


0-82-0-83 




f 


0-89-0-91 


blandula 


m 


0-51-0-67 




f 


0-59-0-72 


burckhardti 


m 


0-62-0-63 




f 


0-64-0-65 


cataloniensis 


m 


0-61-0-68 




f 


0-64-0-73 


cognata 


m 
f 


0-80 
0-83 


genistae 


m 
f 


0-72 
0-74-0-76 


hodkinsoni 


m 


0-64-0-69 




f 


0-69-0-74 


horvathi 


m 


0-69-0-79 




f 


0-71-0-83 


klapperichi 


m 
f 


0-45 
0-49 


lautereri 


m 


— 




f 


0-69 


lusitanica 


m 


0-62-0-65 




f 


0-64-0-67 


maculipennis 


m 

f 


0-57-0-68 
0-63-0-68 


magna 


m 
f 


0-89-0-98 
0-99-1-02 


maura 


m 






f 


0-58-0-65 


nervosa 


m 


0-74-0-81 




f 


0-73-0-86 


nigralineata 


m 
f 


0-74 
0-81 


poggii 


m 
f 


0-54-0-59 
0-58-0-63 


pseudoretamae 


m 
f 


0-65-0-66 
0-71-0-73 


pyrenaea 


m 
f 


0-62-0-78 
0-71-0-85 



1-47-1-66 2-25-2-50 — 

1-55-1-69 2-28-2-48 0-59-0-63 

1-32-1-37 1-84-1-91 

1-25-1-34 2-15-2-24 0-54-0-59 

1-63-1-67 1-91-1-95 

1-65-1-68 2-48-2-50 0-82-0-84 

1-14-1-32 1-70-2-01 

1-22-1-40 1-70-2-30 0-56-0-62 

1-14-1-16 1-86-1-90 — 

1-13-1-16 2-18-2-20 

1-18-1-34 1-69-2-04 

1-19-1-36 1-96-2-33 0-67-0-74 

1-63 2-10 

1-76 2-46 0-93 

1-21 1-99 

1-22-1-24 2-16-2-33 0-87-0-93 

1-42-1-54 1-90-2-00 — 

1-52-1-68 2-26-2-38 0-71-0-79 

1-80-2-04 2-51-2-73 — 

1-77-2-04 2-68-3-06 0-75-0-89 

— 1-50 

1-86 0-60 

1-91 — 

1-63 2-12 0-71 

1-17 0-67-0-85 

1-09-1-24 1-84-1-97 0-53-0-55 

1-25-1-40 1-84-2-13 — 

1-16-1-38 2-03-2-24 0-54-0-59 

2-33-2-56 3-09-3-35 

2-48-2-67 3-60-3-77 1-04-1-08 
unknown 

1-23 1-81-2-27 0-55-0-58 

1-88-2-06 2-39-2-72 

1-80-2-18 2-60-3-15 1-09-1-15 

1-76 2-33 — 

1-97 2-71 0-86 

1-06-1-41 1-33-1-57 — 

1-06-1-19 1-76-1-94 0-58-0-59 

1-38-1-40 1-91-1-99 

1-40-1-42 2-18-2-29 0-60-0-61 

1-50-1-72 2-02-2-49 

1-54-1-82 2-33-2-85 0-76-0-83 



0-37-0-39 0-35-0-37 0-26-0-28 

0-35-0-38 0-33-0-36 0-28-0-30 

0-40-0-41 0-31-0-33 0-31-0-33 

0-29-0-34 0-22-0-26 0-19-0-23 

0-32 0-29 0-29 

0-30-0-34 0-25-0-29 0-24-0-26 

0-38 0-35 0-31 

0-40 0-42 0-36 

0-32-0-33 0-32-0-33 0-28-0-29 

0-40-0-47 0-29-0-35 0-28-0-29 

0-24 0-23 0-20 

0-28 0-29 0-25 

0-29-0-32 0-24-0-26 0-17-0-21 

0-32-0-34 0-27-0-31 0-22-0-23 

0-44-0-50 0-44-0-47 0-38-0-40 

male unknown 

0-45-0-49 0-39-0-41 0-35-0-39 

0-38 0-31 0-30 

0-27-0-29 0-26-0-27 0-19-0-20 

0-31-0-36 0-24-0-25 0-22-0-23 

0-37-0-41 0-26-0-31 0-25-0-29 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



41 



Table 3 - cont . 













Female 


Male 


Male 


Male 






Head 


Antennal 


Forewing 


proctiger 


proctiger 


paramere 


aedeagus 






width 


length 


length 


length 


length 


length 


length 


radiata 


m 


0-73-0-82 


1-59-1-78 


2-70-2-80 


_ 


0-40-0-41 


0-35-0-37 


0-30 




f 


0-78-0-84 


1-82-1-92 


2-60-2-70 


0-85 


— 


— 


— 


retamae 


m 


0-84-1-00 


2-19-2-75 


2-81-3-18 


— 


0-46-0-53 


0-43-0-47 


0-37-0-40 




f 


0-95-1-09 


2-21-2-75 


2-96-3-51 


1-00-1-08 


— 


— 


— 


siciliensis 


m 


0-74-0-81 


1-72-1-78 


2-35-2-50 


— 


0-37-0-40 


0-37-0-39 


0-31-0-32 




f 


0-80-0-85 


1-86-1-88 


2-69-2-79 


0-83-0-87 


— 


— 


— 


spartiisuga 


m 


0-73-0-75 


1-81-1-83 


2-43-2-44 


— 


0-36-0-38 


0-31-0-33 


0-33-0-35 




f 


0-81-0-82 


1-94-1-96 


2-80-2-86 


0-94 


— 


— 


— 


spectabilis 


m 


0-94-1-00 


2-80-3-06 


3-42-3-52 


— 


0-51-0-54 


0-41-0-44 


0-35-0-37 




f 


1-00-1-09 


3-29-3-52 


3-83-4-14 


0-90-0-92 


— 


— 


— 


syriaca 


m 


0-67-0-70 


1-67-1-88 


1-95-1-97 


— 


0-35-0-42 


0-30-0-34 


0-24-0-27 




f 


0-65-0-78 


1-89-1-90 


2-17-2-30 


0-67-0-69 


— 


— 


— 


ulicis 


m 


0-70-0-75 


1-37-1-50 


1-69-1-81 


— 


0-36-0-37 


0-35-0-36 


0-28-0-29 




f 


0-74-0-79 


1-38-1-60 


1-84-2-10 


0-78-0-85 


— 


— 


— 


variegata 


m 


0-67-0-78 


1-78-2-23 


2-76-3-06 


— 


0-40-0-44 


0-30-0-34 


0-26-0-29 




f 


0-71-0-87 


1-74-2-25 


2-82-3-59 


0-82-0-84 


— 


— 


— 


vicina 


m 


0-82-0-83 


1-41-1-50 


2-12-2-20 


— 


0-36-0-45 


0-38-0-44 


0-33-0-35 




f 


0-86-0-87 


1-39-1-42 


2-37-2-39 


0-77-0-83 


— 


— 


— 


vittipennella 


m 


0-71-0-76 


1-74-1-81 


2-35-2-58 


— 


0-37-0-42 


0-37-0-40 


0-30-0-33 




f 


0-67-0-75 


1-48-1-79 


2-42-2-66 


0-80-0-81 


— 


— 


— 



ARYTAINILLA Loginova 

Spartina Heslop-Harrison, 1951: 443; 1961a: 417. Type-species: Psylla spartii Guerin, by monotypy. 
[Homonym of Spartina Harris & Burrows, 1891.] 

Lindbergia Heslop-Harrison, 1951: fig. 2, a,b. Nomen nudum [no included species] {nee Lindbergia 
Riedel, 1958). 

Arytainilla Ramirez Gomez, 1956: 76. Nomen nudum [type-species not designated]. 

Alloeoneura (Hispaniola) Ramirez Gomez, 1956: 91, partim. Nomen nudum [type-species not desig- 
nated]. 

Arytainilla Loginova, 1972a: 17. Type-species: Psylla delarbrei Puton, 1873, designated by Loginova, 
1972a: 17. 

Comments. All described species of Arytainilla occur in the west Mediterranean area and the Macarone- 
sian Islands (Table 4). This is an heterogeneous genus and is almost certainly not a monophyletic group. 
The type-species, A. delarbrei, is exceptional in having 8-segmented antennae, long genal cones, an oval 
forewing with a short pterostigma and unusual male parameres. Most other congeners have 10-segmented 
antennae, short genal cones, an oblong-oval forewing and simple parameres. Arytainilla species are well 
illustrated by Sulc (1907a; 1910a, b) and Loginova (1972a; 1916b) and there is little point in repeating their 
descriptions. Further collecting is required to establish relationships between the species and we have 
confined ourselves to recognising species-groups where possible (Table 4). The spartiophila-group is 
separated from the rest of Arytainilla by the characteristic upturned and pointed apex of the female 
proctiger. The dividens-group comprises species that have oblong-oval forewings, short genal cones, 
moderate to long female terminalia of normal form and male parameres of various shape but most usually 
slender and with a bulge or projection on the anterior margin. The proboscidea-subgroup is defined by the 
elongate rostral segments and the spotted patterning along the forewing veins. Members of the dividens- 
subgroup have parameres with a large bulge on the anterior margin, whereas those in the prognata- 
subgroup have a smaller ill-defined bulge. The equitans-subgroup consists of species similar to those above 
but with parameres that lack the anterior bulge. 

The remaining species tend to have unique characters that set them apart from the rest of the genus. A. 
delarbrei has already been mentioned; it does, however, have a tendency to the same form of female 
terminalia as that of the spartiophila-group. A. cytisi has a truncate female ventral valve and a massive 
ovipositor. A. umbonata has a more elongate, parallel-sided forewing and a distinct bulge on the ventral 
valve of the female terminalia. The female of A egena has long genal cones and short terminalia that are of 



42 



D. HODKINSON & D. HOLLIS 



a characteristic shape. Arytainilla nubivaga (Loginova) comb. n. and A. devia (Loginova) comb, n., two 
distinctive Macaronesian species, are transferred to Arytainilla from Arytaina until a more satisfactory 
position can be found for them within the classification. Nevertheless, they do show some similarity 
with delarbrei and the remaining species, A. ima, particularly in the longer genal cones and the fore wing 
shape of A. devia. The paramere shape in these two Canary Island species is complex and unique in the 
genus. 



Table 4 Species-groups within Arytainilla with information on host plant and distribution. 



Host plant 



Distribution 



spartiophila-group 

spartiophila (Forster, 1848) 



spartiicola (Sulc, 1907) 

gredi (Ramirez Gomez, 1956) comb. n. 

dividens-group 
dividens-subgroup 
dividens Loginova, 1976b 
modica Loginova, 1976b 
hakani Loginova, 1972a 
cognata Loginova, 1972a 
pileolata Loginova, 1976b 
proboscidea-subgroup 

proboscidea Loginova, 1976b 
nigralineata Loginova, 1976b 

program-subgroup 

prognata Loginova, 1976b 

incuba Loginova, 1976b 
equitans-subgxoxxp 

equitans Loginova, 1976b 

diluta, Loginova, 1976b 

Residual species 

delarbrei (Puton, 1873) 

cytisi (Puton, 1876) 



ima Loginova, 1972a 
egena Loginova, 1972a 
umbonata Loginova, 1976b 
devia (Loginova, 1976b) comb. n. 

nubivaga (Loginova, 1976b) comb. n. 



Cytisus scoparius 


Britain 

Spain 

France 




Netherlands 




Switzerland 




Austria 


Cytisus scoparius 


Portugal 

France 

Switzerland 


Cytisus scoparius? 


Spain 


Cytisus proliferus 
unknown 


Canaries 
Canaries 


unknown 


Morocco 


unknown 


Morocco 


unknown 


Canaries 


Adenocarpus anagyrus 
Adenocarpus or 


Canaries 
Canaries 


Cytisus 




unknown 


Canaries 


unknown 


Madeira 


unknown 


Canaries 


unknown 


Canaries 


Genista tinctoria 


France 


Calicotome spinosa 


Spain 

Algeria 

France 


unknown 


Israel (IDH) 

Italy 

Spain 

Yugoslavia 

Turkey 

Morocco 


Retama raetam 
indet. legume 
Cytisus proliferus 


Morocco, Libya (BMNH) 
Canaries 


Adenocarpus anagyrus 
Cytisus fragrans 


Canaries 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 43 

Discussion 

The legume-feeding psyllid fauna of the west Palaearctic appears to comprise a number of 
evolutionary lines that are identified and diagnosed in this paper as separate monophyletic 
genera or species-groups within the Acizziinae and Arytaininae. Arytaina is defined by 
characters that include the broad form of the male paramere, the aedeagus with a strongly 
hooked apex and the lack of a pterostigma and costal break in the forewing. Livilla is defined by 
the narrow elongate paramere and the long bulbous or weakly hooked apex of the aedeagus. 
Cyamophila possesses a paramere that resembles that of Arytaina but the propleurites/ 
pronotum structure is more advanced and the costal break and pterostigma are well-developed 
in the forewing. We have suggested that Arytainilla is heterogeneous and probably comprises a 
number of distinct evolutionary lines. Acizzia is separated from the other genera, into the 
subfamily Acizziinae, by the presence of lateral lobes on the male proctiger. The extent of 
homoplasy in several of the other characters examined makes it difficult to suggest clear 
relationships between genera. Thus we have concluded that Arytaina, Livilla, Cyamophila and 
Arytainilla, together with the monotypic central Asian genus Astragalita, should be retained, for 
the present, in the subfamily Arytaininae. We recognise, however, that there are no good 
autapomorphies that define the subfamily and separate it from the Psyllinae. It remains, 
nevertheless, a convenient and widely recognised grouping. The Acizziinae is a distinct 
evolutionary line warranting subfamily status. 

Arytaina, Livilla and some groups within Arytainilla are most diverse in the Mediterranean 
Basin and are associated with papilionoid legumes of the tribe Genisteae, which itself has a 
primarily Mediterranean distribution. By contrast, most Cyamophila species occur in the arid 
regions of central Asia and are associated with other tribes of papilionoid legumes, particularly 
Galegeae, Trifoleae, Loteae and Sophoreae. The genus is not known from Africa and the one 
west European species, C. prohaskai, represents the western limit of this genus. Acizzia species 
are widespread throughout Africa, Arabia, India, South East Asia and Australasia. They are 
primarily associated with the equally widely distributed mimosoid legume genera Acacia and 
Albizia. Acizzia appears to be an old genus with a very wide distribution that reaches its natural 
northern and western limits in the area under consideration. 

Legume systematics, particularly of the tribe Genisteae, have been the subject of much 
debate (Bisby, 1981). It is of interest to see if the pattern of host plant relationships for the 
different psyllid groups can contribute to an understanding of legume classification and vice 
versa. In a recent classification of the legumes Bisby (1981) split the Genisteae into two 
subtribes, the Lupininae and the Genistinae. He subdivided the latter into three main groups of 
genera (Table 5). His Cytisus-group contains the arytainine host genera Cytisus, Laburnum and 
Chamaecytisus; the Genista-group contains Genista, Ulex and Retama, while the outliers include 
Adenocarpus, Calicotome and Spartium. 

Table 5 shows the distribution of psyllids across their host plants in the Genisteae. Each record 
indicates a known association between a psyllid species and a species of host plant. Species of 
Livilla are concentrated on hosts within Bisby's Genista-group, particularly the large genus 
Genista itself, but also on Ulex and Retama. However, some species also feed on genera in 
Bisby's other two groups of Genistinae. Of the two psyllid species known to feed on the outlier 
group, spectabilis is monophagous on Spartium junceum, whereas pyrenaea is recorded from 
Calicotome and Genista. Among the three species feeding on the Cy r/sws-group {radiata, 
cognata and variegata) cognata also has a known host in the Genista-group, while radiata is 
oligophagous on several species of Chamaecytisus. The core species of Livilla, the spartiisuga- 
and horvathi-groups, tend to occur on Genista. The morphologically isolated species, such as 
variegata and spectabilis, tend to occur on the more taxonomically isolated groups of host plants. 

By contrast with Livilla, the hosts of Arytaina and Arytainilla are more evenly spread across 
the Genisteae. Within the Cytisus-group, Arytaina genistae and A. maculata occur on Cytisus 
and Chamaecytisus, whereas Arytainilla species such as gredi, spartiophila, spartiicola, devia, 
dividens and nubivaga are concentrated on Cytisus. The Genista-group supports just four 
species, Arytaina genistae (record requires confirmation), A. torifrons and Arytainilla delarbrei 



44 



D. HODKINSON & D. HOLLIS 



Table 5 Host plant relationships of Livilla, Arytaina and Arytainilla species across the Genisteae. The 
number of species in each genus (N) is taken from Bisby (1981) except for Argyrolobium which is taken 
from Flora Europaea (Tutin et al. , 1964). 









Number of host records 






N 


Livilla 


Arytaina 


Arytainilla 


Total 


Subtribe Lupininae 
Subtribe Genistinae 
















Cytisus-group 
Laburnum 


2 


2 








2 


Hesperolaburnum 
Podocytisus 
Cytisophyllum 
Petteria 




























Argyrocytisus 
Cytisus 


33 




2 




1 



6 



9 


Chaemaecytisus 
Genista-group 
Retama 


30 
4 


6 

3 


5 



2 
1 


13 

4 


Genista 


87 


18 


2 


1 


21 


Echinospartum 
Stauracanthus 


3 

2 


















Ulex 


20 


2 








2 


Outlier group 
Argyrolobium 
Adenocarpus 
Calicotome 


2 
15 

2 




1 




2 
1 




2 
1 



4 

3 


Erinacea 


1 














Spartium 
Gonocytisus 


1 

3 


1 












1 




on Genista, and Arytainilla egena on Retama. The outlier plant group supports Arytaina putonii 
and Arytainilla cytisi on Calicotome, and Arytaina adenocarpi and Arytainilla devia on Adeno- 
carpus. 

Table 5 indicates that the larger the plant genus the more psyllid species that genus is likely to 
support, with a psyllid feeding record for about every four species of plants. This relationship can 
be interpreted in two ways. It may indicate a simultaneous diversification of both plants and 
insects (coevolution), but, more likely, it is the result of an initial plant species radiation with a 
subsequent radiation of the insects, made successful by their high degree of monophagy (see 
Benson et al. , 1976; Vane-Wright, 1978). 

Overall, however, our present level of knowledge of psyllid host plant relationships adds little 
to an understanding of the classification and evolution of the Genisteae. There is already a high 
level of incongruence between legume classifications based on morphology and those based on 
other biological characteristics (Bisby, 1981). The lack of clear specialisations by the different 
psyllid groups adds further to this incongruity. 

References 

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Andrianova, N. S. & Klimaszewski, S. M. 1983. A contribution to the knowledge of psyllids (Psylloidea) in 
southern and south eastern European U.S.S.R. Acta Biologica, Katowice 13: 30-46. 

Arzone, A. & Vidano, C. 1985. II fitomizo Psylla uncatoides su mimosa in Liguria. Informatore Fitopatalo- 
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Aulmann, G. 1912. Beitrage zur Kenntnis der Psylliden Fauna von Neu-Guinea. Entomologische Rund- 
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1913. Psyllidarum Catalogus, 92pp. Berlin. 

Baeva, V. G. & Kankina, V. K. 1971. The Psyllids (Homoptera: Psylloidea) of Pamir. Izvestiya Acaedemii 
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W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 45 

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Psyllidae). Lancashire and Cheshire Naturalist 8: 387-390. 
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ve zirai Karantina genel mudurlugu, Izmir bolge zirai miicadele arastirma entitusii mudurlugu, Arastirma 

Eserleri Serisi 44: 1-122. 
Oshanin, B. 1907. Verzeichnis der Palaarktishen Hemipteren. Band 2 Homoptera part 2. Ezhegodnik 

Zoologcheskogo Muzeya 12: 338-381. 
1912. Katalog der Palaarktischen Hemipteren (Heteroptera, Homoptera- Auchenorhyncha und Psyl- 
loidea), xvi + 187 pp. Berlin. 
Ossiannilsson, F. 1952. Catalogus insectorum Sueciae XII. Hemiptera, Homoptera, Psylloidea. Opuscula 

Entomologica 17: 193-200. 

1963. Notes on British psyllids (Hem., Horn.). Entomologist 96: 249-257. 

1971. Till Kannedomen om Kullabergs Halvvingar (Hemiptera). Kullabergs Natur 14: 1-55. 

Pettey, F. W. 1924. South African psyllids. Entomology Memoirs, Department of Agriculture, Union of 

South Africa 2: 21-30. 

1925. New South African psyllids. South African Journal of Natural History 5: 125-142. 

1933. New species of South African Psyllids 3. Entomology Memoirs, Department of Agriculture, 

Union of South Africa 8: 3-23. 
Pflugfelder, O. 1941. Psyllina. In Bronns, H. G. (Ed.), Klassen und Ordnungen des Tierreichs. 4: part 3, 

book 8. Leipzig. 
Preisner, H. 1927. Eine neue Psylla aus den Ostalpen (P. prohasakai). Konowia 6: 263-266. 
Prohaska, K. 1928. Beitrag zur Kenntnis der Psylliden (Blattflohle) Karntens. Zeitschrift des Osterreichis- 

chen Entomologen-Vereins 13: 4-6. 
Puton, A. 1871. Description de deux nouvelles especes de Psyllides. Annales de la Societe Entomologique 

de France 1:435-439. 
1873. Notes pour servir a l'etude des Hemipteres. Annales de la Societe Entomologique de France 3: 

11-26. 
1876. Notes pour servir a l'etudes des Hemipteres. Description des especes nouvelles ou peu connues. 

Annales de la Societe Entomologique de France 6: 275-290. 
1878. Psylla (Arytaina) retamae Puton. Bulletin de la Societe Entomologique de France 1878: 

CXXXIV-CXXXV. 

1886. Catalogue des Hemipteres de la faune Palearctique, 3rd edn, 99 pp. Caen. 

1899. Catalogue des Hemipteres de la faune palearctique, 4th edn, 121 pp. Caen. 



Ragusa, E. 1907. Emmitteri di Sicilia. II Naturaliste siciliano 19: 209-237. 

Ramirez Gomez, C. 1956. Los Psilidos de Espana. B olefin de la Real Sociedad Espanola de His tor ia Natural 

(Biologica) 54: 63-106. 
Rapisarda, C. 1985. Sulla presenza in Italia di Acizzia acaciaeb alley anae (Froggatt) (Homoptera, 

Psylloidea), nuovo parassita di Acacie ornamentali. Informatore Fitopatologico 10: 45-49. 
Reiber, F. & Puton, A. 1880. Catalogue des Hemipteres-Homopteres de l'Alsace et de la Lorraine et 

supplement au catalogue de Hemipteres-Heteropteres. Bulletin de la Societe d'Histoire Naturelle de 

Colmar 20-21: 49-77. 
Reuter, O. M. 1875. Heteropterorum novorum species aliquot descripsit. Notiser ur Sails kapets pro Fauna 

et Flora Fennica Forhandlingar , 11: 328-333. 
1881. Till Kannedomen om sveriges Psylloder. Entomologisk Tidskriftl: 145-172. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 51 

1909. Charakteristik und Entwickelungsgeschichte der Hemipteren-Fauna (Heteroptera, 

Auchenorryncha und Psyllidae) der palaarktischen Coniferen. Acta Societatis Scientiarum Fennicae 

36: 1-129. 
Rudow, F. 1875. Zur Kenntnis der Psylloden Norddeutschlands. Programm der Realschule in Neustadt- 

Eberwelde 1875: 3-14. 
Samy, O. 1973. Psyllids of Egypt [Homoptera: Psyllidae]. Bulletin de la Societe Entomologique d' Egypte 

56: 437-480. 
Schaefer, H. A. 1949a. Beitrage zur Kenntnis der Psylliden der Schweiz. Mitteilungen der Schweizerischen 

Entomologischen Gesellschaft 22: 1-96. 
1949b. Biologische und okologische Beobachtungen an Psylliden (Hemiptera). Verhandlungen der 

Naturforschenden Gesellschaft in Basel 60: 25-41. 
Schwarz, E. A. 1904. Notes on North American Psyllidae. Proceedings of the Entomological Society of 

Washington 6: 234-245. 
Scott, J. 1876. Monograph of the British species belonging to the Hemiptera - Homoptera, family 

Psyllidae. Transactions of the Entomological Society of London 24: 525-569. 
1879. Description of a new species of the family Psyllidae - Floria horvathi n. sp. Entomologist's 

Monthly Magazine 16: 84-85. 
1880. Description of the nymph of Arytaina genistae Latr. Entomologist's Monthly Magazine 17: 

132-133. 
1882a. Food plants and times of appearance of the species of Psyllidae found in Great Britain together 

with others which may be expected to occur here. Entomologist's Monthly Magazine 19: 13-15. 
1882/?. The British Psyllina, with corrections in the synonymy. Entomologist's Monthly Magazine 18: 

253-256. 
Smreczynski, S. 1954. Materialy do fauny pluskwiakow Polski 3. Psylloidea. Fragmenta Faunistica 7: 

Sulc, K. 1905. Revise Psyll sbirky Dudovy. Casopis Ceskoslovenske Spolecnosti Entomologicke 2: 1-4. 
1907. Prispevky ku poznani Psyll. 1. Psylla spartii Guerin-Loew a Psylla spartiicola n. sp. Rozpravy 

Ceske Akademie Ved, Praha (section 2) 16: 1-8. 
1910a. Uvod do studia, synoptika tabulka a synonymicky katalog druhu rodu Psylla, palaearcticke 

oblasti. Sitzungsberichte der K. Bohmischen Gesellschaft der Wissenschaften 2 (1909): 1-46. 

1910b. Prispevky k poznani II. Rozpravy Ceske Akademie Ved, Praha (section 2) 19: 1-32. 

Szulczewski, J. W. 1927. Materialien zur Psyllidfauna Grosspolens. Sprawozdania Komisji Fizyograficznej 

Oraz Materialy do Fiziografii Kraju 61: 197-204. 
Tamanini, L. 1955. Alcuni nuovi repert di Psyllidi Italiani e Francesci (Homoptera, Psyllina). Bolletino 

della Societa Entomologica Italiana 85: 10-11. 
1977. Notizie corologiche e morphologiche su alcuni psyllidi poco noti delle prealpi (Homoptera: 

Psylloidea). Studi Trentini di Scienze Naturali 54: 103-119. 
Thomson, C. C. 1877. XXIX. Ofversigt af Skandinaviens Chermes-arter. Opuscula Entomologica (Editit 

C. G. Thomson), Trelleborg 8: 820-841. 
Tut hill, L. D. 1943. The Psyllids of North America north of Mexico (Psyllidae: Homptera) (Subfamilies 

Psyllinae and Triozinae). Iowa State College Journal of Science 17: 443-660. 

1952. On the Psyllidae of New Zealand. Pacific Science 6: 83-125. 

1964. Homoptera, Psyllidae. Insects of Micronesia 6: 353-376. 

Tutin, T. G. etal. (Eds) 1964-1980. Flora Europaea. Cambridge. 

Van der Goot, P. 1912. Naamlist van inlandsche Psyllidae. Entomologische Berichten, Amsterdam 3: 

281-285. 
Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north of Mexico excepting the Aphididae, 

Coccidae and Aleurodidae. University of California Publications, Technical Bulletin!'. 1-902. 
Vane-Wright, R. I. 1978. Ecological and behavioural origins of diversity in butterflies. In Mound, L. A. & 

Waloff, N. (Eds), Diversity of insect faunas. Symposia of the Royal Entomological Society of London 9: 

56-70. 
Vondracek, K. 1951a. A new Psyllid from Tanger. Casopis Moravskeho Musea v Brne 36: 119-122. 
1951/?. Jumping plant-lice in the collections of the Moravian Museum (Brno). Melichar's collection 

revised, part 1 . Casopis Moravskeho Musea v Brne 36: 123-131. 
1952. Results of zoological scientific expedition of the National Museum in Praha to Turkey. Sbnornik 

Entomologickeho Oddeleni Ndrodniho Musea v Praze 28: 435-450. 

1953. Jumping plant-lice in the collections of the Moravian Museum (Brno). Casopis Moravskeho 



Musea v Brne 38: 174-179. 
1957. Mery Psylloidea. Fauna CSR 9: 1-431 



52 I. D. HODKINSON & D. HOLLIS 

Wagner, W. & Franz, H. 1961. Uberfamilie Sternorrhyncha (Psylloidea), pp. 158-179. In Franz, H. (Ed.), 

Die Nordst-Alpen im Spiegal ihrer Landtierwelt 2: 729 pp. 
Wahlgren, E. 1934. Svenska bladdlopper (Chermesidae). Entomologisk Tidskrift 55: 81-104. 
Waltl, J. 1837. Bemerkungen iiber einige Insekten. Isis, Leipzig 1837: 277-279. 
Watmough, R. H. 1968a. Population studies on two species of Psyllidae (Homoptera: Sternorrhyncha) on 

broom (Sarothamnus scoparius (L.) Wimmer). Journal of Animal Ecology 37: 283-314. 
1968ft. Notes on the biology of Arytaina spartiophila Forster and A. genistae Latreille (Homoptera: 

Psyllidae) on broom {Sarothamnus scoparius (L.) Wimmer) in Britain. Journal of the Entomological 

Society of Southern Africa 31: 115-122. 
White, I. M. & Hodkinson, I. D. 1982. Psylloidea (Nymphal Stages). Handbook for the Identification of 

British Insects 2 (5b): 1-50. 
1985. Nymphal taxonomy and systematics of the Psylloidea. Bulletin of the British Museum (Natural 

History) (Entomology) 50: 153-301. 
Yang, C. T. 1984. Psyllidae of Taiwan. Taiwan Museum Special Publications Series 3: 1-205. 
Yen, A. 1977. Redescription of the species of Psylla Geoffroy (Hemiptera: Psyllidae) originally described 

by Froggatt. Journal of the Australian Entomological Society 16: 7-20. 
Zgardinska, A. 1976. Koliszki (Homoptera: Psyllidae) Okolic Siedlec. Polski Pismo Entomologiczne 46: 

241-246. 






W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



53 










Ml? 



Figs 1-8 Acizzia species. 1,2, bicolorata: (1) forewing; (2) head. 3, 4, hollisi: (3) forewing; (4) head. 5 6 
uncatoides: (5) forewing; (6) head. 7, 8, acaciaebaileyanae: (7) forewing; (8) head. 



54 



D. HODKINSON & D. HOLLIS 







W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



55 




fi^'fL % am °P hlla s P ecies ' 21 - 2 ^ glycyrrhizae: (21) forewing; (22) head; (23) cf genitalia, lateral 
m?A a 2 ^ amere ', inn f F V,eW; (25) apical Segment of aedea g"S. 26-30, pro/^a/: (26) forewing; 
(27) head; (28) cf genitalia, lateral view; (29) cf paramere, inner view; (30) apical segment of aedeagus 



56 



I. D. HODKINSON & D. HOLLIS 




,*, 31-42 Cy^cpUI. **» 31-36, »«..« (Ilo Hi™» «* tJ^'SSlw 9 

(4™5 paramere; (41) apical segment of aedeagus; (42) $ termmaha, lateral view. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



57 




R m^ k 50 ^ ^[T SpedeS - 43 ' 44 ' lon Z icella - ( 43 > brewing; (44) head. 45, 46, ge/i»tee: (45) forewing- 
fofeJin (50) head "^ (4?) forewin g' ( 48 ) head - 49 > 5 °> maculata (Spanish material): (49) 



58 



D. HODKINSON & D. HOLLIS 




-^.iTS.&S^S^^^i^A^'SS'^ 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



59 




F 1orewTnt r^7S TUT' ^ ■?' ^T^ (59) f0reWing; (60) heacL 61 ' 62 ' ad ™ocarpi: (61) 
torewing, (62) head. 63, 64, magmdentata; (63) forewing; (64) head. 



60 



I. D. HODKINSON & D. HOLLIS 




Figs 65-74 Arytaina species. 65-67, teniae ^ ^^l^f^^r^Z^J 
view; (67) apical segment of aed eagus- 68-70; "^fijg > ° f flM (Sim ish material); (71) cT 
paramere, inner view; (70) apical segment of aedeagus_ 71 ,i macuu v £ ?4 fricam . $ 

genitalia, lateral view; (72) Cf paramere, inner view; (73) apical segment ot aeaeag 
genitalia, lateral view (after Heslop-Harnson). 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



61 




^f" 84 Arytama species. 75-77, putonii: (75) cf genitalia, lateral view; (76) tf paramere, inner view; 
(77) apical segment of aedeagus. 78-80, torifrons: (78) cf genitalia, lateral view; (79) a* paramere, inner 
JZSfii ' api .° a segment of aed eagus. 81, angustatipennis: cf genitalia, lateral view; 82-84, W 
(82) CT genitalia, lateral view; (83) cf paramere, inner view; (84) apical segment of aedeagus 



mnica: 



62 



D. HODKINSON & D. HOLLIS 




Figs 85-94 Arytaina and Cyamophila species. 85-87, A. adenocarpi: (85) tf genitalia, lateral view; (86) 
tf paramere'inner view; (87) apical segment of aedeagus. 88-90, A magnidentata. f)<*J^»> 
lateral view; (89) cT paramere, inner view; (90) apical segment of aedeagus. 91 A. atlasiensis V 
terminalia, lateral view. 92, A. hispanica: ? terminalia, lateral view. 93, C. g/ycyrr/nzae: $ terminal*, 
lateral view. 94, C. prohaskai: $ terminalia, lateral view. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



63 




Figs 95-102 Arytaina and Livilla $ terminalia, lateral view. 95, A. longicella. 96 A venistae 97 A 
maculata. 98, L. radiata. 99, A. adenocarpi. 100, A. putonii. 101, A ™A* W^ 



64 



D. HODKINSON & D. HOLLIS 




109 



(110) head. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



65 




117 



Figs 111-118 Livilla species. Ill, 112, Aiervo^: (111) forewing; (112) head. 113, 114, mgra/mea/a: (113) 
fo[eZg-(118)h^ M: ° 15) f ° reWing; (116) head ' H7 ' H8 ' vit «P ennella - ( 117 ) 



66 



I. D. HODKINSON & D. HOLLIS 




126 



Figs 119-126 Livilla species. 119, 120, retamae: (119) forewing; (120) head 121 122, auteren: (121) 
forewing; (122) head 123, 124, syriaca; (123) forewing; (124) head. 125, 126, radiata; (125) forewing; 
(126) head. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



67 




127 




129 




131 





133 




134 



¥ Tu?Unlt L/V mm P K de J- JS" 1 £ c ^~f ( 127 ) dewing; (128) head. 129, 130, maculipennis: 
fore^a^i S (B1) f ° reWing; (B2) hCad - 133 ' 134 '^^ fl 33 ) 



68 



D. HODKINSON & D. HOLLIS 




142 



forewing; (142) cf forewing; (143) head. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



69 




144 





146 




Fi n4^hi 5 H iff ^ eCieS ' 144 7^' b < iandula - (144) for ewingof lectotype; (145) forewing (Morocco)- 
head lusitamca: (147) forewing; (148) head. 149, 150, bimaculata: (149) forewing; (150) 



70 



D. HODKINSON & D. HOLLIS 




153 




155 




157 




forewing; (I 58 ) head - 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



71 




166 



^ore^n^r^T '£?' m adUSt r (159) f ° rewin ^ ( 160 > head - 161 ' 162 > spectabilis, (161) 
Sngio^held" Vaf ^ flte: (163) f ° reWing; (164) hCad - 165 ' 166 ' *"**■"«: (165) 



72 



D. HODKINSON & D. HOLLIS 




Figs 167-178 Livilla species. 167-169, ulicis: (167) cf genitalia, lateral view; (168) cf paramere outer 
view (169) apical segment of aedeagus. 170-172, bivittata: (170) cf genitalia, lateral view; (171) Cf 
paramere, outer view; (172) apical segment of aedeagus. 173-175, vicina: (173) Cf genitalia lateral 
view; (174) cT paramere, outer view; (175) apical segment of aedeagus. 176-178, cognata: (1/0) O 
genitalia lateral view; (177) cf paramere, outer view; (178) apical segment of aedeagus. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



73 




187 



Figs 179-190 Livilla species. 179-181, nervosa'. (179) O" genitalia, lateral view; (180) d" paramere, outer 
view; (181) apical segment of aedeagus. 182-184, nigralineata: (182) cf genitalia, lateral view (183) O* 
paramere, outer view; (184) apical segment of aedeagus. 185-187, horvathi: (185) cf genitalia, lateral 
view; (186) cf paramere, outer view; (187) apical segment of aedeagus. 188-190, vittipennella: (188) <$ 
genitalia, lateral view; (189) c? paramere, outer view; (190) apical segment of aedeagus 



74 



D. HODKINSON & D. HOLLIS 




202 



Figs 191-202 Livilla species. 191-193, retamae: (191) C? genitalia, lateral view; (192) cf paramere, outer 
view; (193) apical segment of aedeagus. 194-196, lautereri: (194) cf genitalia, lateral view; (195) cf 
paramere, outer view; (196) apical segment of aedeagus. 197-199, syriaca: (197) cf genitalia, lateral 
view; (198) cf paramere, outer view; (199) apical segment of aedeagus. 200-202, radiata: (200) cf 
genitalia, lateral view; (201) cf paramere, outer view; (202) apical segment of aedeagus. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



75 



207 




211 



Figs 203-214 Livilla species. 203-205, cataloniensis: (203) tf genitalia lateral view (204) cf mramere 

outer v^w; (205) ap 1C al segment of aedeagus. 206-208, macllipennis: (206 ^ g"n ta^i fatera Tew-' 

207) cTparamere, ^outer view; (208) apical segment of aedeagus .7^-211 t sLaielsis^^T^Z: 

O genitalia, lateral view, (213) tf paramere, outer view; (214) apical segment of aedeagus. 



76 



I. D. HODKINSON & D. HOLLIS 



219 




225 



223 



Fies 215-226 Livilla species. 215-217, magna: (215) Cf genitalia, lateral view; (216) C? paramere outer 
Wewf(2n) apical segment of ^^^mfl^^tf^^^ffl 
paramere, outer view; (220) apical segment of aedeagus. m-™Mdk™om£K\W &^*>™™* 
view (222) d 1 paramere, outer view; (223) apical segment of aedeagus. 224-226, poggu. (224) O 
genitalia, lateral view; (225) tf paramere, outer view; (226) apical segment of aedeagus. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



77 




231 



235 



238 



v ew mv W ' SPeC1 ' S - f 7 "i 29 ' gmiStae: (227) tf S enitalia ' lateral vi ™; (228) tf paramere outer 
SSSff^^^T^ 230 " 2 , 32 ' "«"** ( 23 °) <* genitalia, lateral view; (231)^ 
later7vi™ C' ( } ^^ Segment of aedea g us - 233-235, pseudoretamae: (233) tf genitalia 

55X ffite^y 1 " view; (235) apical segment of aedeagus - 236 - 238 ' *^« S 

O genitalia, lateral view, (237) cf paramere, outer view; (238) apical segment of aedeagus. 



78 



i. D. HODKINSON & D. HOLLIS 







249 



247 



Figs 239-250 Livilla species. 239-241, adusta: (239) tf genitalia, lateral view; (240) d" paramere outer 
itorOAVt apical segment of aedeagus. 242-244, spectabilis: (242 cf genitalia, lateral view (243) tf 
752£ SSS vS^SU) apical sfgment of aedeagus. 245-247, ^^i^^S^ 
view; (246) cT paramere, outer view; (247) apical segment of aedeagus. 248-250 burckhardu. (248) O 
genitalia, lateral view; (249) cT paramere, outer view; (250) apical segment of aedeagus. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 




253 




^ei^L^ewl?^ AdZZia and C y am °P hila s P edes - 251-253, L. lusUamca: (251) a* 
genitalia, lateral view, (252) cT paramere, outer vew; (253) apical segment of aedeaa.is ?S4 ?^ / 

sSsK 8 ^ iatera : r w; (255) * p ara - re - o^?^rs?ffiii 2 S2i of 

aedeagus. 257 258, Arytama afncana (after Heslop-Harrison): (257) forewing- (258) head 255 S! 



80 



I. D. HODKINSON & D. HOLLIS 




Figs 263-270 LiviUa, $ terminal lateral view. 263, ulicis. 264, Mnttata. 265, vicina. 266, cognata. 267, 
adjusta. 268, spectabilis. 269, variegata. 270, bwckhardtt. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



81 




Figs 271-278 Livilla, $ terminalia, lateral view. 271, cataloniensis. 272, maculipennis. 273, siciliensis 
274,pyrenaea. 275, magna. 276, spartiisuga. 277, hodkinsoni. 278, blandula. 



82 



I. D. HODKINSON & D. HOLLIS 




285 



Figs 279-286 Livilla, ? terminalia, lateral view. 279, lusitanica. 280, bimaculata. 281 , poggii. 282, maura. 
283,genistae. 284, klapperichi. 285, pseudoretamae. 286, syriaca. 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



83 




Figs 287-294 Livilla ! and ilcteu, 2 terminalia, lateral view. 287, L. nervosa. 288, L. nigralienata 289 



84 



Acacia 2, 3,4,5,43 
Acacia bailey ana 5 

confusa 6 

floribunda 6 

koa6 

koaia 6 

podalyriaefoliae 5 

raddiana 5, 6 

saligna 6 

verniciflua 6 
Acacieae 4 

Adenocarpus 42, 43, 44 
Adenocarpus anagyrus 42 

complicates 13 

hispanicus 13 
A/fctzw 2, 3, 4, 43 
Albizia lophantha 6 
Anthyllis 8 
Anthyllis vulneraria 9 
Argyrocytisus 44 
Argyrolobium 44 
Astragalus 8 
Astragalus sp. 10 

Calicotome 43, 44 
Calicotome spinosa 18, 22, 34, 42 
Chamaecytisus 43, 44 
Chamaecytisus austriacus 15, 27, 
29 

borysthenicus 17, 29 

heuffelii 15 

ratisbonensis 17, 26, 29 

supinus 26 
Cytisophyllum 44 
Cyrims 42,43,44 
Cy tows-group 43, 44 



I. D. HODKINSON & D. HOLLIS 

Index to host plants 

Cytisus fragrans 42 
proliferus 42 
purgans 39 
scoparius 2, 15, 39, 42 

Echinospartium AA 
Erinacea AA 

Galegeae 2, 7, 43 
Gemsta 43, 44 
Ge/mta-group 43, 44 
Genista acanthoclada 31, 33 

aetnensis 23, 36 

Corsica 22, 37 

fasselata 21, 31 

germanica 26 

hirsuta23, 35 

hispanica 19 

pZ/osa 22, 34 

radwta25,27,28 

salzmanii 34 

spp. 16, 25, 32, 33, 39 

tinctoria 15, 2A, 42 

triacanthos 23 , 36 
Genisteae 2, 3, 11,19,43,44 
Genistinae 43, 44 
Glycyrrhiza 8 
Glycyrrhiza glabra 8 
Gonocytisus AA 

Hesperolaburnum AA 

Ingae 4 

Laburnum 43, 44 



Laburnum alpinum 38 

anagyroides 38 

spp. 23 
Leguminosae 7 
Lemboptropis nigricans 29 
Loteae 2, 7, 43 
Lupininae 43, 44 

Medicago 8 
Medicago sativa 9 

Ononis spinosa 24 

Petteria AA 
Podocytisus AA 

RetamaA3,AA 

Retama monosperma 30 

raeta/n 30, 42 

sp.21 

sphaerocarpa 30 
Rosaceae 7 

Sophoreae 2, 7, 43 
Spflrfwra43,44 
Spartium junceum 21, 37, 43 
Stauracanthus AA 

Trifoleae 2, 7, 43 

Me* 43, 44 

£//e* europaeus 24 
parviflorus 33 

V/cw 8 
View sp. 9 



Index 

Invalid names are in italics; principal references are in bold 



acaciae (Psylla) 4 
acaciaebaileyanae (Acizzia) 4 

(key), 5, 6, 10 
acaciaebaileyanae (Psylla) 5 
acaciaebaileyanae-group 4 (key), 

5 
acaciae-baileyanae (Arytaina) 5 
acaciae-baileyanae (Psylla) 5 
Acizzia 2, 3 (key), 4, 10, 43 
Acizziinae 2, 3 (key), 4, 43 
adenocarpi (Arytaena) 13 
adenocarpi (Arytaina) 12, 13 

(key), 16, 17, 18,44 
adusta {Florid) 39 
adusta (Livilla) 21 (key), 39, 40 
africana 12 (key), 13-14, 18 
albizziae (Acizzia) 11 
albizziae (Arytaina) 11 
Allaeoneura 19 



Alloeoneura 2, 19, 29, 41 
alpina {Florid) 38 
Amblyrhina2, 10, 11,19,26 
Amblyrrhina 25 
Amorphicola 2 

angustatipennis {Amblyrhina) 14 
angustatipennis (Arytaina) 12 

(key), 14, 18 
annosa {Floria (Brinckitia)) 19 
antennata (Retroacizzia) 11 
Arylaina 15 
Arytaena 15, 24 
Arytaina 2, 3 (key), 7, 10-11, 17, 

18,24,42,43,44 
Arytainilla 2, 4 (key), 41, 42, 43, 

44 
Arytaininae 2, 3 (key), 4, 7, 43 
Arytainini 2, 7 
assimilis (Arytaina) 11 



assimilis (Ceanothia) 11 
Astragalita 2, 43 
Ataenia 15 
atlasiensis 12 (key), 14, 18 

bicolor (Arytaina) 1 1 
bicolor (Ceanothia) 11 
bicolorata (Acizzia) 5 (key), 7, 10 
bicolorata (Psylla) 7 
bimaculata 22 (key), 34-35, 40 
bivittata 3, 4, 20 (key), 24, 25, 40 
blandula {Floria) 35 
blandula (Livilla) 23 (key), 35, 36, 

40 
boharti (Arytaina) 11 
boharti (Ceanothia) 11 
brevigena (Arytaina) 11 
brevigena (Insnesia) 11 
Brinckitia 19 



W. PALAEARCTIC LEGUME-FEEDING PSYLLIDS 



85 



burckhardti 3, 20 (key), 25, 40 
burckhardti-group 19, 20 (key), 25 

callunae (Psylla) 23 
cataloniensis {Florid) 31 
cataloniensis (Livilla) 22 (key), 

32-33, 40 
Ceanothia 11 
Chermes 15 
Ciriacreminae 2 
cognata Low {Amblyrhina) 25 
cognata Low {Amblyrrhina) 25 
cognata Loginova (Arytainilla) 42 
cognata (Low) (Livilla) 3, 20 

(key), 25-26, 40, 42, 43 
coleoptera (Psylla) 23 
cornicola (Arytaina) 1 1 
crawfordi (Insnesia) 11 
Cyamophila 2, 3 (key), 7, 9, 10, 43 
Cyamophilini 2 
cytisi Becker (Psyllodes) 29 
cytisi (Puton) (Arytainilla) 41, 42, 

44 

delarbrei (Arytainilla) 41, 42, 43 

delarbrei (Psylla) 41 

devia (Arytaina) 11 

devia (Arytainilla) 4, 11, 42, 43, 44 

diluta 42 

dividens 42, 43 

dividens-group 41, 42 

dividens-subgroup 41, 42 

egena41,42, 44 
equitans 42 

equitans-subgroup 41, 42 
essigi (Arytaina) 1 1 
essigi (Ceanothia) 11 
Euphalerus 11 
Euryconus 11 

fabra (Psylla) 7 
fasciata (Arytaina) 11 
flava (Arytaina) 11 
flava (Insnesia) 11 
Floria 2, 19 
Floriella 19 

genistae Fabricius (Psylla) 15 
genistae (Latreille) (Arylaina) 15 
genistae (Latreille) (Arytaena) 15 
genistae (Latreille) (Arytaina) 2, 

12 (key), 14-15, 16, 17, 18,43 
genistae (Latreille) (Chermes) 15 
genistae (Latreille) {Chermes 

(Ataenia)) 15 
genistae Latreille (Psylla) 10, 14 
genistae Ramirez Gomez (Livilla) 

3, 20 (key), 38-39, 40 
glycyrrhizae (Cacopsylla) 8 
glycyrrhizae (Cyamophila) 8 

(key), 10 
glycyrrhizae (Psylla) 8 
glycyrrhizae (Psyllodes) 8 
gredi 42, 43 



hakani 42 

hispanica 12 (key), 16, 18 

Hispaniola 41 

hodkinsoni (Floria) 33 

hodkinsoni (Livilla) 22 (key), 33, 

40 
hollisi5(key),6, 10 
hollisi-group 4 (key), 6 
horvathi (Floria) 26 
horvathi (Livilla) 21 (key), 26-27, 

40 
horvathi-group 19, 20 (key), 26, 

27, 39, 43 

ima42 
incuba 42 
Insnesia 11 
insolita (Arytaina) 11 
insolita (Ceanothia) 11 
iolani (Arytaina) 11 
iolani (Insnesia) 11 
isitis (Euphalerus) 11 
isitis (Psylla) 11 

klapperichi 21 (key), 28-29, 40 

lactea (Psylla) 29 

lautereri 21 (key), 30-31, 40 

Lindbergia 41 

lineata (Floria) 28 

Livilla 2, 4 (key), 19,26,29,35, 

39,40,43,44 
longicellall(key),16, 18 
lusitanica 23 (key), 35, 36, 40 

maculata (Amblyrhina) 16 
maculata (Amblyrrhina) 16 
maculata (Arytaina) 12 (key), 14, 

15,16-17,18,43 
maculipennis 22 (key), 32, 33-34, 

40 
maculosa (Amblyrhina) 16 
magna Hodkinson & Hollis 

(Livilla) 23 (key), 36, 40 
magna Crawford (Psyllopa) 10, 15 
magnidentata 13 (key), 17, 18 
maura Loginova (Floria) 35 
maura Vondracek (Floria) 39, 40 
maura Vondracek (Livilla) 22, 23 

(key), 35, 39-40 
medicaginis (Cacopsylla) 9 
medicaginis (Cyamophila) 8 (key), 

9,10 
medicaginis (Psylla) 9 
meridionalis (Arytaina) 11 
meridionalis (Insnesia) 11 
mitella (Arytaina) 11 
mitella (Ceanothia) 11 
modica 42 

montana (Arytaena) 24 
montana (Arytaina) 24 
mopanei (Arytaina) 11 
mopanei (Retroacizzia) 11 

Neopsylla 4 

nervosa 20 (key), 27, 40 



nigralineata Loginova 

(Arytainilla) 42 
nigralineata Hodkinson & Hollis 

(Livilla) 21 (key), 27, 40 
nubivaga (Arytaina) 11 
nubivaga (Arytainilla) 4,11, 42, 

43 

obscura (Acizzia) 11 
obscura (Psyllopa) 11 
odontopyx 8 

pechai (Amblyrhina) 11 

pechai (Spanioneura) 1 1 

Peripsyllopsis 11 

pileolata 42 

poggii (Floria) 36 

poggii (Livilla) 22 (key), 36-37, 40 

proboscidea 42 

proboscidea-subgroup 41 , 42 

prognata 42 

prognata-subgroup 41, 42 

prohaskai (Cyamophila) 8 (key), 

9, 10, 43 
prohaskai (Psylla) 9 
Pseudacanthopsylla 2 
pseudoretamae 21 (key), 35-36, 

40 
Psylla 11, 15, 19,23 
Psyllidae 2 
Psyllinae 2, 4, 7, 43 
Psyllopa 10 
pulchra Crawford, 1919 

(Arytaina) 11 
pulchra (Crawford, 1919) 

(Euryconus) 11 
pulchra Crawford, 1920 

(Arytaina) 11 
pulchra (Crawford, 1920) 

(Insnesia) 11 
punctinervis (Arytaina) 11 
punctinervis (Euphalerus) 1 1 
putoni 17 

putonii (Amblyrhina) 17 
putonii (Arytaina) 12 (key), 

17-18,19,44 
pyrenaea (Floria) 34 
pyrenaea (Floria (Floriella)) 34 
pyrenaea (Livilla) 22 (key), 34, 40, 

43 
pyrenaea (Psylla) 19, 34 
pyrenea 34 

radiata (Aleuroneura) 29 
radiata (Allaeoneura) 29 
radiata (Alloeoneura) 29 
radiata (Arytaina) 19, 29 
radiata (Livilla) 20 (key), 29, 41, 

43 
radiata (Psylla) 29 
radiata-group 19, 20-21 (key), 27, 

29 
ramarkrishni 11 
retamae (Alloeoneura) 30 
retamae (Floria) 30 
retamae (Floria (Brinckitia)) 30 



86 



D. HODKINSON & D. HOLLIS 



retamae (Livilla) 21 (key), 30, 31, 

36,41 
retamae (Psylla) 30 
Retroacizzia 11 

siciliensis 22 (key), 32, 36, 41 
Spanioneura 11 
spartii Guerin (Psylla) 41 
spartii (Hartig) (Arytaina) 15 
spartii Hartig (Psylla) 10, 15 
spartiicola42,43 
spartiisuga {Florid) 31 
spartiisuga (Livilla) 22 (key), 

31-32, 41 
spartiisuga (Psylla) 31 
spartiisuga-group 19, 21-22 (key), 

31,39,43 
Spartina 41 

spartiophila (Arytainilla) 2, 42, 43 
spartiophila (Psylla) 18 
spartiophila-group 41, 42 
spectabilis {Florid) 37 
spectabilis (Livilla) 20, 21 (key), 

37,41,43 



spectabilis (Psylla) 37 
spinosa ( Acizzia) 1 1 
spinosa (Arytaina) 1 1 
stocklosai8(key), 10 
syriaca {Florid) 31 
syriaca (Livilla) 21 (key), 31, 41 
syriacea 31 

thakrei (Arytaina) 11 
thakrei (Insnesia) 11 
torifrons {Amblryhina) 18 
torifrons (Arytaina) 12 (key), 16, 

18-19, 43 
torifrons (Psylla) 10, 18 
tuberculata (Arytaina) 11 
tuberculata (Insnesia) 11 
turkiana {Amblyrhina) 11 
turkiana (Spanioneura) 11 

uichancoi (Arytaina) 1 1 
uichancoi (Insnesia) 11 
ulicis (Curtis, 1835) (Arytaina) 15 
ulicis Curtis, 1835 (Psylla) 15 
ulicis (Curtis, 1835) {Arytaend) 15 



ulicis Curtis, 1836 (Livilla) 4, 19, 
20 (key), 23-24, 25, 41 

ulicis-group 19, 20 (key), 23, 39 

umbonata41, 42 

uncata (Psylla) 5 

uncatoides (Acizzia) 4 (key), 5-6, 
10 

uncatoides (Psylla) 5 

unctata 5 

variabilis (Arytaina) 11 
variabilis (Insnesia) 11 
variegata {Florid) 38 
variegata (Livilla) 22, 23 (key), 38, 

41,43 
vicina {Florid) 24 
vicina (Livilla) 3, 4, 20 (key), 

24-25, 41 
virgina 11 
vitipennella 28 
vittipennella {Floria) 26, 28 
vittipennella (Livilla) 21 (key), 28, 

41 
vittipennella (Psylla) 28 



British Museum (Natural History) 

Milkweed butterflies: their cladistics and biology 

P. R. Ackery & R. I. Vane- Wright 

The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of 
their biology have stimulated far more attention than can be justified by species numbers 
alone. In recent years, an expansive literature has grown, considering aspects of their 
courtship and pre-courtship behaviour, migration, larval hostplan associations, mimicry and 
genetics. The popularity of danaines among biologists can certainly be attributed to this 
combination, within one small group, of so many of the factors that make butterflies such an 
interesting group to study. The obvious need to place this wealth of biological data within an 
acceptable systematic framework provided the impetus for this volume. 

Started eight years ago within the conventions of evolution by natural selection and 
Hennig's phylogenetic systematics, the book is now largely about natural history (what the 
animals have and do, where they live and how they develop) and natural groups - as revealed 
by a form of analysis approaching that practised by the new school of 'transformed cladistics'. 
The authors have prepared a handbook that will appeal to a wide range of biologists, from 
museum taxonomists to field ecologists. 

425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. 
ISBN 565 00893 5. 1984. Price £50. 



Titles to be published in Volume 56 



The legume-feeding psyllids (Homoptera) of the west Palaearctic Region 

By I. D. Hodkinson & D. Hollis 

A review of the Mai vales-feeding psyllid family Carsaridae (Homoptera) 

By D. Hollis 

The world genera of Rhadalinae (Coleoptera: Melyridae) 

By E. R. Peacock 



Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk 
Printed in Great Britain by Henry Ling Ltd, Dorchester 



Bullet i of the 

British Museum (Natural History) 



A review of the Malvales-feeding psyllid 
family Carsidaridae (Homoptera) 

D. Hollis 



BRITISH MUSEUM 
(NATURAL HiSTORYl 

-2 NOV 1987 

PRESENTED 
ENTOMOLOGY LI6RARY 



Entomology series 
Vol 56 No 2 



29 October 1987 



The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, 
and an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
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specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
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Parts are published at irregular intervals as they become ready, each is complete in itself, 
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London SW7 5BD, 
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) 



© British Museum (Natural History), 1987 



The Entomology series is produced under the editorship of the 

Keeper of Entomology: Laurence A. Mound 

Publications Manager (Entomology): W. Gerald Tremewan 






ISBN 565 06029 5 
ISSN 0524-6431 



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British Museum (Natural History) 
Cromwell Road 
London SW7 5BD 



Entomology series 
Vol 56 No 2 pp 87-127 



Issued 29 October 1987 




A review of the Malvales-feeding psyllid family 
Carsidaridae (Homoptera) 



David Hollis ^o^ og* 



Department of Entomology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Contents 

Synopsis 87 

Introduction 87 

Material, methods and terminology 88 

Carsidaridae Crawford 89 

Key to genera 90 

Epicarsa Crawford 91 

Carsidara Walker 92 

Protyora Kieffer 94 

Tyora Walker 95 

Tenaphalara Kuwayama 97 

Allocarsidara gen. n 100 

Paracarsidara Heslop-Harrison 104 

Mesohomotoma Kuwayama 106 

Phylogeny , biogeography and hostplants 108 

List of characters 109 

References 112 

Hostplant index 126 

Index 126 

Synopsis 

The psyllid family Carsidaridae is characterised and considered to be the sister-group of the Homotomi- 
dae. It is shown that the monophyletic group, Carsidaridae + Homotomidae, is trophically linked to 
hostplants of the dicotyledonous superorder Malviflorae. Eight genera, Epicarsa, Carsidara, Tyora, 
Tenaphalara, Protyora, Paracarsidara, Mesohomotoma and a new genus Allocarsidara, are recognised in 
the family; they are diagnosed and a key is given for their identification. It is considered that Mastigimas is 
not a carsidarid and is provisionally placed in the Calophyidae. Strogylocephala is confirmed as a good 
genus, with Synaphalara syn. n. as its junior synonym; it is removed from the family and placed, again 
provisionally, in the Calophyidae. Species within each genus are reviewed; six new species are described, 
eight new combinations are proposed and one subspecies is given species status. Hostplants of all 
recognised species are restricted to the families Sterculiaceae, Bombacaceae and Malvaceae, within the 
order Malvales. A phylogeny for the Carsidaridae is proposed which suggests that the group followed the 
radiation of the Malvales prior to the separation of the African and South American continents. 

Introduction 

The Psylloidea or jumping plant-lice comprise a group of small, phloem-feeding sternorrhyn- 
chous Homoptera with a world-wide distribution. Individual psyllid species show a high degree 
of hostplant specificity, particularly during the larval stages, and related species tend to develop 
on related species or groups of dicotyledonous angiosperms. At present about 2000 species are 
described but collecting in the last 20 years in the tropics and southern temperate regions 
indicates that the group is probably four times as large. 

Until recently the higher classification of the group was based on the Holarctic fauna but 
this has proved inadequate for the inclusion of the rest of the world's species. With such a 



v: 



Bull. Br. Mm. nat. Hist. (Ent.) 56 (2): 87-127 Issued 29 October 1987 



55 D. HOLLIS 

large proportion of undescribed taxa it is useful to be able to classify them into biologically 
meaningful groups and use their high degree of hostplant specificity to predict probable host 
ranges. 

White & Hodkinson (1985) attempted, with a good measure of success, to produce such a 
classification based largely on larval morphology and cladistic methodology. It is now desirable 
to test this classification using adult morphology. White & Hodkinson's Clade 2 (Family 
Triozidae) has been reviewed (Hollis, 1984) and shown to be a good monophyletic group. They 
suggested that the sister-group of the Triozidae is the family Carsidaridae (Clade 4) and I 
attempt here to test the monophyly of this group, discuss its sister-group relationships, diagnose 
the included genera and review the known species. 

Carsidarids are restricted in their hostplant preferences to the families Sterculiaceae, Bomba- 
caceae and Malvaceae within the Malvales, although three recently described Chinese species 
(Yang & Li, 19856) have been recorded (possibly as vagrants) from other, unrelated families. 
Each genus that is recognised here is usually restricted to only one of the above families or to the 
Malvaceae and Bombacaceae together, but the Mesohomotoma species-complex seems able to 
develop on Malvaceae and Sterculiaceae. Several of the host species are of some economic 
importance, e.g. cocoa {Theobroma cacao), durian {Durio zibethinus), silk cotton (Bombax 
buonopozense and B. ceiba) and kapok (Ceiba pentandra); several strong-fibre sources (Abuti- 
lon indicum, Firmiana simplex and Hibiscus tiliaceus); and Heritiera littoralis, the wood of which 
is used in boat-building. Psyllid damage to these plants is not well-documented but high 
infestations of Tenaphalara acutipennis on Bombax ceiba cause wilting and premature leaf-fall 
(Sohi & Sandu, 1978), and Mesohomotoma tessmanni can cause desiccation and death of 
terminal shoots of Theobroma cacao in Africa (Wood, 1980). 

Life history information is available for Tenaphalara acutipennis on Bombax ceiba (= 
malabaricum) in Bihar, India (Rahman, 1932), Mesohomotoma tessmanni on cocoa in Ghana 
(Kaufmann, 1973) and Protyora sterculiae on Br achy chiton populneum in Australia (Froggatt, 
1923; Morgan, 1984). In these species the larvae are free-living, tend to feed in aggregations and 
produce copious amounts of wax during their later stages. Larvae are known for a further nine 
species and all are morphologically similar, suggesting similar lifestyles. 

Little is known of the natural enemies of the group. Cotterell (1927) recorded larvae of the 
syrphid flies Baccha sinuata, B. praeusta and B. picta feeding on colonies of larvae of 
Mesohomotoma tessmanni in West Africa, and Leston & Gibbs (1968) recorded the mirid bug 
Deraeocoris crigi preying on the same psyllid. Bhatia & Shaffi (1933) recorded the larvae of the 
syrphid flies Baccha pulchriformis and Sphaerophoria javana feeding on larvae of Tenaphalara 
acutipennis on Bombax ceiba in India. 

Materials, methods and terminology 

Most of the material studied is deposited in the British Museum (Natural History) (BMNH), 
with supplementary specimens from the Bernice P. Bishop Museum, Honolulu (BPBM), and 
the National Museum of Natural History, Washington (USNM). Type material is deposited in 
BMNH; BPBM; USNM; Australian National Insect Collection, Canberra (ANIC); Beijing 
Agricultural University Insect Collection, China (BAUIC); Forest Research Institute, Dehra 
Dun, India (FRI); Institut Royal des Sciences Naturelles de Belgique, Bruxelles (IRSNB); 
Museum fur Naturkunde der Humboldt-Universitat, Berlin (MNHU); National Collection of 
Insects, Plant Protection Research Institute, Pretoria (NCI); Naturhistorisches Museum, 
Vienna (NM); Osaka Museum of Natural History, Japan (OMNH); and Termeszettudomanyi 
Muzeum, Budapest (TM). 

All measurements are quoted in millimetres and were taken from slide-mounted specimens. 
Reference points, from which measurements were taken of the various structures, are given in 
Hollis (1976; 1984). 

Apart from the head illustrations, all figures were drawn from slide-mounted specimens; 
setosity of heads and forewings is not shown. The inner surface of the right paramere is shown, 
apart from Figs 47, 49, 52, 55, which were drawn in anterior view. Dorsal and ventral views of the 



MALVALES-FEEDING PSYLLIDS 89 

caudal areas of larvae are illustrated. Structural terminology follows that of Vondracek (1957) 
and Hollis (1976; 1984). 

Acknowledgements 

I am grateful to the following colleagues for providing loans and gifts of material, and further 
information: Dr D. R. Miller and Miss Louise Russell, United States Department of Agricul- 
ture, Systematic Entomology Laboratory, Beltsville; Keith Arakaki and Wayne Gagne, BPBM; 
J. Etienne, formerly of Institut Senegalaise Recherches Agricoles, Ziguinchor; and Dr T. 
Vasarhelyi, Termeszettutomanyi Miizeum, Budapest. Peter Broomfield, Entomology Depart- 
ment, BMNH, contributed considerably with the head and forewing illustrations, and other 
colleagues in the Department offered valuable criticisms of earlier drafts of the paper. Special 
thanks are due to the staff of the Systematic Entomology Laboratory, USDA, Beltsville, for 
their companionship and accommodation during my stay in Washington. 

CARSIDARIDAE Crawford 

Prionocnemidae Scott, 1882: 466. [Invalid name.] 

Prionocnemina Scott; Low, 1886: 160. 

Prionocneminae Scott; Froggatt, 1901: 286; Kieffer, 1906: 387. 

Carsidarinae Crawford, 1911: 481, in part; 1914: 53, in part; 1919: 155, in part; Wu, 1935: 126; Tuthill, 

1950: 52, in part; Brues et al. , 1954: 156; Heslop-Harrison, 1958: 578, in part; Braza & Calilung, 1981: 

352, in part; Yang, 1984: 168, in part; Brown, 1985: 245. 
Tenaphalarini Heslop-Harrison, 1958: 578, in part. 
Carsidaridae Crawford; Klimaszewski, 1964: 12, fig. 7, in part; Bekker-Migdisova, 1973: 98, in part; 

Morgan, 1984: 118, in part; Brown, 1985: 244, in part. 

Diagnosis. Antennal sockets enlarged and swollen ventromedially and vertex often deeply divided by 
median suture giving the head a 'cleft' appearance in dorsal view; antennal flagellum with a single subapical 
rhinarium present on flagellomere 3 in addition to those on flagellomeres 2, 4, 6 and 7. False (non- 
tracheate) rs-m crossvein present in forewing or Rs and M 1+2 in broad contact; costal break absent (present 
in Protyora); radular spinules sharply defined into narrow triangular patches, often absent from cu } , a 
patch of spinules present in cell c+sc. Ventral sense organs of hind femur in a basal position and not 
arranged linearly; hind tibia with a well-developed basal spine; hind basitarsus with a single apical spur. 
Male subgenital plate with a pair of secondary lobes anterior to parameres, these lobes appearing to be 
sclerotised projections arising from the membrane lining the inner surface of the subgenital plate. 

Final instar larva elongate, clearly divided into head, thorax and abdomen; antennae elongate, 
10-segmented; legs elongate, tarsal arolium sessile and fan-shaped or globular; wing buds small, without 
humeral lobes; thoracic sclerites poorly differentiated; caudal region of abdomen differentiated and 
bearing convoluted pore bands, anus terminal or terminodorsal; body setae mainly simple but scattered, 
small, lanceolate setae present on caudal sclerites marginally and submarginally. 

Hostplants. Malvales (Sterculiaceae, Bombacaceae and Malvaceae). 

Discussion. An historical account of the early conceptual development of the family is given by Hollis 
(1976: 4). Heslop-Harrison (1958) considered the group as a tribe within the Ciriacreminae including the 
genera Carsidara, Mastigimas, Mesohomotoma, Protyora, Epicarsa and Diceraopsylla, and separated 
these genera from another of his Ciriacremine tribes, the Tenaphalarini, containing Tenaphalara, 
Strogylocephala, Togepsylla and Tyora. Hollis (1984) referred Dicer aopsylla to the Aphalaridae and White 
& Hodkinson (1985) placed Togepsylla in the same family. 

Vondracek (1957: fig. 52) considered the Carsidaridae as the sister-group of the Triozidae, differentiat- 
ing both families from the Ciriacreminae which he placed in the Psyllidae. Klimaszewski (1964: fig. 7) 
agreed with these concepts. Bekker-Migdisova (1973) recognised seven subfamilies in her interpretation of 
the Carsidaridae which amounted to a very large assemblage of many quite unrelated genera. 

White & Hodkinson (1985: 242, 273) considerably restricted their concept of the family to the genera 
Mastigimas, Tenaphalara, Protyora, Mesohomotoma, Paracarsidara, Carsidara and tentatively Aconop- 
sylla, Carsidaroida, Nesiope and Tyora. They placed Epicarsa in the Phacopteronidae, considering the 
anastomosis of Rs and M 1+2 as the important synapomorphy. For reasons given below (p. 90) Mastigimas 
is now removed from the family and placed provisionally in the Calophyidae; Carsidaroida and Nesiope are 
synonyms of Tyora, and Aconopsylla is referred to the Aphalaridae, close to Haplaphalara and Diclidoph- 



90 D. HOLLIS 

lebia which are also Malvales-feeding. Brown (1985) removed Epicarsa from the Phacopteronidae to the 
Carsidaridae, using the presence of supplementary lobes on the male subgenital plate as a more reliable 
synapomorphy. This character, and the presence of a non-tracheate rs-m crossvein in the forewing, are the 
best pair of autapomorphies for the Carsidaridae. A crossvein of this type is unique to the family. The 
supplementary male subgenital lobes occur elsewhere in the Psylloidea only in the Oriental Cedrela- 
feeding genus Trisetipsylla Yang & Li (1985a) which, in other characters, is a member of the family 
Psyllidae. The male hypovalves and tracheate rs-m crossvein of Ciriacremum (Psyllidae) are not homo- 
logues (Hollis, 1976). 

White & Hodkinson diagnosed the family on the following larval characters: anal pore field arranged in 
convoluted bands; dorsal surface of abdomen lacking distinct sclerites; body margin and dorsal surface 
without sectasetae or derivable structures. To this character-set can be added that poorly defined 
lanceolate setae are present marginally and submarginally on the caudal plate. These same authors 
postulated that the Carsidaridae formed the sister-group of the Triozidae, citing the absence of a costal 
break and the presence of fewer than six apical spurs on the hind tibia as adult synapomorphies, but they 
suggested no larval synapomorphies. These two adult characters are probably unreliable as they are 
subject to homoplasy within the Psylloidea generally. 

Brown (1985) considered the family to be the sister-group of the F/cus-feeding Homotomidae, citing the 
'cleft head', weakly sclerotised A 2 vein and the absence of Aj in the hindwing and a patch of spinules in 
c+sc of the forewing as synapomorphies . This is a more acceptable hypothesis , although Aj is present in the 
hindwing of many homotomids. Further synapomorphies for the two groups are the development of a pair 
of strong epiphyses on the metapostnotum, and the position and arrangement of the ventral sense organs of 
the hind femur. The former structures appear to function as 'tethering posts' for both pairs of wings when 
these are in the resting position. Similar but probably non-homologous structures occur on the metas- 
cutellum in some triozids. The basal position of the ventral sense organs on the hind femur is considered to 
be primitive (Hollis, 1984: 11) but in homotomids and carsidarids they are not linearly arranged and the 
proximal sensillum is placed at right-angles to the other two. 

White & Hodkinson (1985: 274) provisionally placed Mastigimas in the Carsidaridae, presumably on 
larval characters, and Brown (1985) did not dispute this. However the only carsidarid adult character which 
Mastigimas displays is the enlarged antennal sockets giving the head a cleft appearance. The apical 
rhinarium is absent from the third flagellomere, there is no rs-m crossvein, no metapostnotal epiphyses, the 
forewing radular areas are diffuse, one of the metafemoral sensoria is medial, the basal spine of the hind 
tibia is absent, there are two apical spurs on the hind basitarsus and there are no supplementary lobes on 
the male subgenital plate. Furthermore the larva lacks lanceolate setae marginally and submarginally on 
the caudal plate, and the host plants of all known species are species of Cedrela (Meliaceae). For these 
reasons Mastigimas is excluded from the Carsidaridae but this raises the difficult question of where to place 
the genus in White & Hodkinson's system. A sister-group relationship cannot be argued for it with either 
the Carsidaridae or the Carsidaridae + Homotomidae and I provisionally place Mastigimas near Apsylla 
which White & Hodkinson tentatively referred to the Calophyidae. 

The Carsidaridae, as it is recognised here, contains eight genera: Epicarsa, Carsidara, Tyora, Tenapha- 
lara, Protyora, Paracarsidara, Mesohomotoma and a new genus, Allocarsidara. These are diagnosed 
below and a key is given for their identification. Nominal species within each genus are discussed in general 
terms. Insufficient material and incomplete hostplant data prevent thorough revision to species level at 
present. 

Key to genera 

1 Forewing widening to a broadly rounded apex, Rs and M 1+2 in broad contact, cuj almost as high 

as wide and with a value of less than 1-3 (Fig. 18); antennal flagellum thickened, 1st flagellar 
segment short and broad, not more than four times longer than its greatest width (Fig. 2) 

EPICARSA (p. 91) 

- Forewing narrowing to a subacute apex , Rs and Mj +2 not in contact but connected by a false rs-m 

crossvein (Figs 19-39), cuj much wider than high and with a value of at least 1-6; antennal 
flagellum elongate, 1st flagellar segment long and narrow, not less than nine times longer than 
its greatest width 2 

2 Pterostigma absent and M+Cu about half as long as Cu stem (Figs 36-39); male proctiger 

bipartite, with a large, anvil-shaped median posterior lobe in addition to lateral lobes (Fig. 77) 

MESOHOMOTOMA(p. 106) 

- Pterostigma present (Figs 19-31 , 35) , if absent (Figs 32-34) then M+Cu almost twice as long as 

Cu stem; male proctiger unipartite, without median posterior lobe, with or without lateral 
lobes(Figs40,43,46,51,57,66,75) 3 



MALVALES-FEEDING PSYLLIDS 91 

3 M+Cu very short, about one-third as long as R stem and less than half as long as Cu stem (Fig. 

19); female terminalia, in profile, rounded dorsally, ventrolateral margins of proctiger with 
dense fringes of setae, lateral palps ridged (Fig. 79) CARSIDARA (p. 92) 

- M+Cu longer, about as long as or longer than R stem and Cu stem (Figs 20-35); female 

terminalia, in profile conical, proctiger sometimes with a median lobe posterior to anal pore, 
lateral palps not ridged (Figs 80-89) 4 

4 False r r rs crossvein present in addition to rs-m crossvein (Figs 20-26) 5 

- rj-rs crossvein absent (Figs 27-35) 6 

5 Radular area present in cu la , claval suture reaching hind margin of forewing very close to apex of 

Cu lb (Figs 20-24) TYORA (p. 95) 

- Radular area absent from cu la , claval suture reaching hind margin of forewing distant from apex 

ofCw /6 (Figs25,26) TENAPHALARA (p. 97) 

6 Costal break present (Figs 27, 28); apical segment of aedeagus with strongly expanded apex 

(Fig. 45); male proctiger without lateral expansions (Fig. 43) PROTYORA (p. 94) 

- Costal break absent (Figs 29-35); apical segment of aedeagus not or only weakly expanded 

apically (Figs 59, 62, 65, 68, 71, 74, 76); male proctiger with lateral expansions (Figs 57, 60, 63, 
66,69,72,75) 7 

7 Anteorbital lobes strongly developed into 'horn-like' structures, lateral and posterior margins of 

vertex rounded (Figs 11, 12); apex of ductus ejaculatorius enclosed within apical lobes of 
aedeagus (Figs 59, 62, 65, 68, 71, 74) ALLOCARSIDARA(p 100) 

- Anteorbital lobes very weak; lateral margins of vertex sharply raised, posterior margin angular 

(Figs 14, 15) ; apex of ductus ejaculatorius extended beyond apical lobes of aedeagus (Fig. 76) 

PARA CARSIDARA (p . 104) 

EPICARSA Crawford 

Epicarsa Crawford, 1911: 488; 1914: 56; Hodkinson & White, 1981: 496; White & Hodkinson, 1985: 241, 
243, 272; Brown, 1985: 258. Type-species: Epicarsa corniculata Crawford, by original designation and 
monotypy. 

Epiciasa Caldwell, 1941: 419. [Misspelling.] 

Epicrasa Caldwell, 1941: 421. [Misspelling.] 

Description (only slide-mounted material available for study). Integument of head and thorax very 
sparsely short setose. Head (Fig. 2) with disc of vertex convex, discal foveae shallow, frontal margin not 
clearly defined but deeply incised by median suture, lateral and hind margins rounded, lateral ocelli placed 
anteriorly on vertex, anterolateral tubercles well-developed; antennal sockets enlarged, flagellum short, 
2-12 times longer than head width, 1st flagellar segment short and swollen, less than 4 times longer than its 
greatest width; genal cones present as a pair of small tubercles on either side of mid ventral suture 
immediately above clypeus; ultimate rostral segment relatively short, less than 3 times longer than its 
greatest width. 

Forewing (Fig. 18) obovate, rounded apically, 2-3-2-6 times longer than wide; costal break absent, 
pterostigma short and triangular, r r rs crossvein absent, Rs arched downwards towards and in broad 
contact with M 1+2 , M+Cu stem slightly longer than Cu stem and about as long as M stem, cuj cell value 
about 1-0 and without radular spinules, apex of claval suture distant from apex of Cu lb ; hindwing with 
grouped costal setae; apical spurs of hind tibia arranged l+2(or 3) + l. 

d" proctiger unipartite with strongly expanded lateral lobes, each with a secondary inwardly pointing 
subapicallobe. 

$ proctiger abruptly stepped in apical fifth, apical part with a dense covering of short, spine-like setae, 
apex upcurved (see Ferris, 1928: figs 3a-c); subgenital plate short, similarly stepped apically. 

Larva. Unknown. 

Hostplant genus. Ceiba (Bombacaceae). 

Comments. Although possessing most of the features of the family character-suite, the forewing shape, 
venation and pattern of Epicarsa (Fig. 18) is quite unlike other carsidarids. These forewing characters are 
similar to many homotomids, the carsidarid sister-group, but the similarity may be convergent. Further 
characters shared between Epicarsa and homotomids are the thickened antennal flagellum (Fig. 2) and the 
bifid lateral lobes of the male proctiger. For these reasons I consider Epicarsa to be the sister-group of the 
rest of the carsidarids. 
Brown (1985: 259, fig. 98H) described the endtube of the ductus ejaculatorius of this genus as being 



92 D. HOLLIS 

barbed and, from this, suggested a close relationship to the other South American carsidarid genus 
Paracarsidara (and therefore, by inference, to Mesohomotoma) . I have not examined this character in the 
holotype of E. corniculata and the only other males known (Ferris, 1928) are whole-mount, permanent, 
microslide preparations in which the structure is obscured. 

Included species 
Epicarsa corniculata Crawford 

(Figs 2, 18) 

Epicarsa corniculata Crawford, 1911: 488; 1914: 56; 19256: 56; Ferris, 1928: 244; Costa Lima, 1942: 103; 

Tuthill, 1950: 58; Hodkinson & White, 1981: 496; Brown, 1985: 260. Holotype cf , Brazil (USNM) [not 

examined]. 
Epiciasa corniculata Caldwell, 1941: 419. [Misspelling.] 
Epicrasa corniculata Caldwell, 1941: 421. [Misspelling.] 

Recorded distribution. Brazil (Crawford, 1911), Panama (Brown, 1985), Mexico (Ferris, 1928). All 
other authors refer to these records. 

Material examined 
2 d\ 2 $, Panama, Mexico (slide-mounted material of Ferris (1928) and Brown (1985)). 

Hostplant. Ceiba sp. 

CARSIDARA Walker 

Carsidara Walker, 1869: 329; Scott, 1882: 466; Kieffer, 1906: 387, in part; Enderlein, 1910: 138; Aulmann, 

1913: 79, in part; Heslop-Harrison, 1960: 241; Hodkinson, 1983: 344; 1986: 303. Type-species: Carsidara 

marginalis Walker, by monotypy. 
Dynopsylla Crawford, 1915: 264, in part. 
Thysanogyna Crawford, 1919: 157; Wu, 1935: 126. Type-species: Dynopsylla minor Crawford, by original 

designation and monotypy. [Synonymised by Crawford, 1924: 619.] 
Eustigmia Enderlein, 1921: 119. Type-species: Dynopsylla minor Crawford, by original designation and 

monotypy. [Junior objective synonym of Thysanogyna.] 

Description. Integument of head and thorax with dense, moderately long setae. Head (Figs 3, 4) with disc 
of vertex concave, discal foveae clearly marked as broad longitudinal oblique grooves, frontal margin 
sharply denned and deeply incised by median suture, lateral and hind margins prominent but obtuse, 
lateral ocelli placed posteriorly on vertex, anterolateral tubercles well-developed; antennal sockets 
enlarged, flagellum 2-4-2-8 times longer than head width, 1st flagellomere elongate, more than 9 times 
longer than its greatest width; genae swollen ventrally on either side of median suture anterior to clypeus, 
each with a prominent papilla, weak lateroventral tubercles also present; apical rostral segment elongate, 
more than four times longer than its greatest width. 

Forewing (Fig. 19) ovate with subacute apex, 2-4-2-9 times longer than wide; costal break absent, 
pterostigma present, r r rs crossvein absent, M+Cu stem very short, about one-third as long as R 
stem and 0-4-0-5 times as long as Cu stem, cuj cell value 2-33-3-75, without radular spinules, apex of claval 
suture adjacent to apex of Cu lb ; hindwing with grouped costal setae; apical spurs of hind tibia arranged 
1+3 + 1. 

C? proctiger (Fig. 40) unipartite, lateral lobes moderately developed; apex of basal segment of aedeagus 
expanded and strongly sclerotised , apical segment of aedeagus highly modified (Fig. 42) , endtube of ductus 
ejaculatorius heavily sclerotised. 

$ proctiger (Fig. 79) without posterodorsal lobe, apex strongly sclerotised and upcurved, posterolateral 
margins with fringes of long setae; lateral palps heavily ridged. 

Larva. Unknown. 

Host plant genera. Brachy chiton and Firmiana (Sterculiaceae). 

Comments. Carsidara is distinct from the rest of the family and is diagnosed by its broader forewing, 
the very short M+Cu stem in the forewing, the modified distal segment of the male aedeagus and the 
structure of the female genital apparatus. I regard the genus as the sister-group of the rest of the family less 
Epicarsa. 



MALVALES-FEEDING PSYLLIDS 93 

Included species 
Carsidara marginalis Walker 

Carsidara marginalis Walker, 1869: 329; Scott, 1882: 467; Aulmann, 1913: 79; Crawford, 1924: 619; 

Heslop-Harrison, 1960: 237 etseq.; Hodkinson, 1983: 344; 1986: 304. Holotype $, 'Celebes' (BMNH) 

[examined]. 
Dynopsylla minor Crawford, 1915: 263. Holotype cf , Philippines (?USNM) [not examined]. [Synony- 

mised by Crawford, 1924: 619.] 
Thysanogyna minor (Crawford) Crawford, 1919: 158; Miyatake, 1981: 37. 
Eustigmia minor (Crawford) Enderlein, 1921: 119. 

Recorded distribution. Philippines, Sulawesi, Tanimbar Is. 

Material examined 

1 Cf , 9 $ , from Malaya, Hong Kong (on Br achy chiton rupestris), Sabah, Sulawesi. 

Hostplant. Br achy chiton rupestris. 

Comments. Crawford (1919) records slight differences in colour and forewing vein proportions in the 
Tanimbar Is specimens. The Malayan and Hong Kong material studied differs from Walker's holotype in 
that there is no brown pattern on the forewing around the radular spinules or apices of M 1+2 , M 3+4 or Cu la , 
and M 1+2 is less strongly arched basally. The significance of these differences is not apparent and further 
material with accurate hostplant data is required. 

Carsidara limbata (Enderlein) 

Thysanogyna limbata Enderlein, 1926: 397; Wu, 1935: 126. Syntypes d\ 9, China [not examined]. 
Carsidara limbata (Enderlein) Hodkinson, 1986: 303. 

Recorded distribution. Only known from the type-series, 'China. Tsingtau'. 

Hostplant. Unknown. 

Comments. Enderlein separated this species from marginalis (as minor) on its larger size (forewing length 
5-25-5-75 mm), the brown pattern on the posterior margin of the forewing and some slight venational 
differences. 

Carsidara shikokuensis (Miyatake) comb, n., stat. n. 

Thysanogyna minor skikokuensis Miyatake, 1981: 37. Holotype d\ Japan: Kashima Is (OMNH) [not 
examined] . 

Recorded distribution. Only known from the type series. 

Hostplants. Firmiana simplex; also found on Mallotus japonicus . 

Comments. Miyatake's figures (1981: figs 1A-G), especially of the male genitalia, show this taxon to be 
quite distinct from marginalis and it is given full species status. However, there is a strong possibility that it 
is synonymous with limbata. 

Carsidara africana sp. n. 

(Figs 3, 4, 19,40-42,79) 

Body ochraceous to mid brown, mesopraescutum with a dark brown anteromedial patch and mesonotum 
dark brown laterally; antennal flagellomeres 1-6 dark brown apically, 7-8 entirely dark brown; forewing 
hyaline, with brown pattern as in Fig. 19. 

Head and dorsum of thorax with moderately long, dense setae. Antennal flagellum 2-7-3-0 times longer 
than head width, 1st flagellomere more than nine times longer than its median width and with 6-12 rhinaria 
in apical third, 2nd flagellomere with 10-20 rhinaria in apical half, 3rd flagellomere with 5-7 rhinaria in 
apical quarter, 4th flagellomere with 2 subapical rhinaria and up to 5 more in apical third, 5th flagellomere 
with or 1 subapical rhinarium and up to 3 more in apical third, 6th flagellomere with 2 subapical rhinaria, 
7th flagellomere with a single subapical rhinarium; ultimate rostral segment more than 6 times longer than 
wide. 

Forewing (Fig. 19) 2-47-2-63 times longer than wide and about 1-6 times longer than hindwing; 



94 D. HOLLIS 

pterostigma short, less than half as long as Rj ; M+ Cu stem about 0-33 times as long as R stem and about 0-5 
times as long as Cu stem, cu 1 value 2-33-3-39. 

Cf genitalia as in Figs 40-42; proctiger about 1-4 times longer than wide; paramere with an outwardly 
directed, rounded, posteroapical lobe in addition to posteroapical hook. 

5 terminalia as in Fig. 79. 

Measurements (3 Q\ 1 $>). Head width, cf 0-95-1-01, 9 1-01; length of antennal flagellum, cf 
2-68-2-82, 2 2-81; length of ultimate rostral segment, cf 0-43-0-45, $ 0-48; length of forewing, cf 
4-98-5-71, £ 5-54; length of hind tibia, cf 1-08-1-24, $ 1-14. 

Larva and hostplant. Unknown. 

Holotype cf , Angola: 7 mis W. Gabela, 16-18. iii. 1972, at light (D. Hollis) (BMNH); dry-mounted. 

Paratypes. Ghana: 2 $, Northern Region, Garu, 250 m, N 10-51-W 0-12, 27.X.1971, Nr 510 soil trap 
(Endrody-Younga). Nigeria: 1 cf , Ile-Ife, 26.ii.1970 (/. T. Medler). Uganda: 2 cf , Kampala, 7. iii. 1933 (H. 
Hargreaves). (BMNH; TM); slide- and dry-mounted. 

Comments. The male and female genitalia of this species are remarkably similar to marginalis but the two 
species differ markedly in antennal structure, that of marginalis being simple, with a single subapical 
rhinarium on flagellomeres 2, 3, 4, 6, and 7. 

Carsidara sp. 

Carsidara sp. Heslop-Harrison, 1960: 241. 

A single damaged female of this genus is in BMNH from Madagascar, Nosy Komba. The antennae are 
missing and I am unable to identify the specimen. 

PROTYORA Kieffer 

Protyora Kieffer, 1906: 390; Aulmann, 1913: 29; Tuthill & Taylor, 1955: 250; White & Hodkinson, 1985: 

242, 273. Type-species: Tyora sterculiae Froggatt, by monotypy. 
Neocarsidara Crawford, 1925a: 32; 1927: 31; Tuthill & Taylor, 1955: 250. Type-species: Tyora sterculiae 

Froggatt, by monotypy. [Junior objective synonym.] 

Description. Integument of head and thorax almost glabrose. Head (Figs 5, 6) with disc of vertex weakly 
concave, foveae present as broad longitudinal grooves, hind and lateral margins of vertex rounded, 
anterior margin not sharply defined but deeply incised by median suture, lateral ocelli placed posteriorly on 
vertex, anterolateral tubercules absent or present; antennal sockets enlarged, antennal flagellum about 3-7 
times longer than head width, 1st flagellomere more than 10 times longer than wide; genal cones present as 
a pair of small tubercles on either side of mid ventral suture immediately anterior to clypeus, lateroventral 
tubercles absent; ultimate rostral segment 2-7-3-8 times longer than wide. 

Forewing (Figs 27, 28) narrow-elongate, 2-9-3-3 times longer than wide, with a narrowly rounded or 
subacute apex, costal break present, pterostigma present, r r rs crossvein absent, M+Cu stem almost as 
long as or longer than R stem and Cu stem, cuj cell value greater than 2-0, radular spinules absent or very 
weakly present in cuj, apex of claval suture distant from apex of Cu lb \ hindwing with grouped costal setae; 
apical spures of hind tibia arranged 1+2+1. 

Cf proctiger (Fig. 43) unipartite, flask-shaped, with poorly developed lateral lobes; apex of aedeagus 
(Fig. 45) with enlarged apical lobes, endtube of ductus ejaculatorius simple. 

$ proctiger (Fig. 80) short, conical in profile, without posterodorsal lobe. 

Larva. Caudal plate as in Fig. 90. 

Hostplant genus. Brachy chiton (Sterculiaceae). 

Comments. The two main features of this genus are the presence of a costal break in the forewing and the 
swollen and modified apex of the aedeagus. The costal break is either primitively retained or secondarily 
derived and, as I consider its absence as a groundplan feature of the family, I postulate the latter. The form 
of the aedeagal apex is also regarded as a derived character but not strictly homologous with that of 
Carsidara. Apart from the simple, flask-shaped male proctiger, Protyora shares other characters, such as 
the narrow-elongate forewing and long M+Cu stem vein, with the Tyora + Tenaphalara group and the 
Allocarsidara + Paracarsidara + Mesohomotoma group; it is regarded as the sister-group of these genera. 
Until now Protyora was thought to be a purely Australian genus, unique in this respect within the 
Carsidaridae. The discovery of another species in Indonesia shows the genus to have a closer relationship 
with the Oriental fauna and to be less isolated. 



MALVALES-FEEDING PSYLLIDS 95 

Tuthill & Taylor (1955) regarded Neocarsidara as a nomen nudum but under Article 12(b)(5) of The 
International Code of Zoological Nomenclature (3rd edn) the name is valid. However, it has the same 
type-species as Protyora and is therefore an objective synonym. 

Included species 

Protyora sterculiae (Froggatt) 

(Figs 27, 90) 

Tyora sterculiae Froggatt, 1901: 289; 1923: 84. Syntypes a\ $ and larvae, Australia (NSW on 
Brachy chiton populneum, QLD on Brachychiton sp.) (ANIC) [not examined]. 

Protyora sterculiae (Froggatt) Kieffer, 1906: 390; Aulmann, 1913: 29; Tuthill & Taylor, 1955: 250; Morgan, 
1984: 37, 68. 

Neocarsidara sterculiae (Froggatt) Crawford, 1925a: 32; 1927: 31. [Invalid combination, Tuthill & Taylor, 
1955:250.] 

Hostplant. Brachychiton populneum (Sterculiaceae). 

Recorded distribution. Australia (QLD, NSW, SA). 

Material examined 
Adults and larvae from Australia (QLD, ACT). 

Comments. Brief accounts of the life-history of this species are given by Froggatt (1923) and Morgan 
(1984). It differs from the new species described below in that the anterolateral tubercles of the vertex are 
poorly developed, the 1st flagellomere is relatively shorter (10 times longer than its greatest width), the 
forewing (Fig. 27) is relatively broader (2-9-3-0 times longer than wide) and has a more rounded apex, the 
pterostigma is broader, M+Cu stem is relatively longer, the male parameres and aedeagal apex are of a 
different shape and the female proctiger is more obtuse apically (Morgan, 1984). 

Protyora wilsonisp. n. 

(Figs 5, 6, 28, 43-45, 80) 

Description. Similar to sterculiae. Differs in that antennal flagellum is 3-28-3-75 times longer than head 
width and has alternating ochraceous and dark brown rings, 1st flagellomere longer and narrower, about 15 
times longer than its greatest width; anterolateral tubercles of vertex more strongly developed. 

Forewing (Fig. 28) 3-14-3-25 times longer than wide and with a more acute apex, pterostigma narrow, 
M+Cu stem almost as long as R stem and Cu stem, cuj cell value 2-35-3-0. 

Cf paramere and aedeagal apex as in Figs 44, 45. 

$ proctiger (Fig. 80) more acute and slightly upcurved apically. 

Measurements (2 cf, 2 $). Maximum width of head, cf 0-70-0-71, $ 0-74-0-77; length of antennal 
flagellum, cf 2-51-2-65, $ 2-52-2-64; length of ultimate rostral segment, cf 0-20, $ 0-21; length of 
forewing, cf 3-67-3-70, $ 4-24-4-28; length of hind tibia, cf 0-62, $ 0-66. 

Larva and hostplant. Unknown. 

Holotype cf , Indonesia: Sulawesi Utara, Dumoga Bone NP, Hogs Back camp, 21. vi. 1985, swept (M. R. 
Wilson) (BMNH); dry-mounted. 

Paratypes. 2 cf, 2 $, same data as holotype; 1 $, 6.vi.l985; 1 Cf, 1 ?, 25.x. 1985 (M. Asche); 1 ?, 
13.xi.-4.xii.1985, malaise trap (BMNH); 2 cf, x.1985 (Bosmans & Van Stalle) (IRSNB). Slide- and 
dry-mounted. 

TYORA Walker 

Tyora Walker, 1869: 330; Scott, 1882: 470; Kieffer, 1906: 387, in part; Enderlein, 1910: 138; Crawford, 
1911: 483; 1920: 355; 1925a: 32; 1927: 30; Aulmann, 1913: 80, in part; Hodkinson, 1983: 366; Yang & Li, 
19856: 206, 215. Type-species: Tyora congrua Walker, by monotypy. 

Nesiope Kirkaldy, 1908: 389; Aulmann, 1913: 81; Crawford, 1919: 160; Tuthill, 1964: 356; Hodkinson, 
1983: 354; 1986: 319. Type-species: Nesiope ornata Kirkaldy, by monotypy. [Synonymised with 
Carsidaroida by Crawford, 1919: 161, and with Tyora by Crawford, 1920: 355.] 

Carsidaroida Crawford, 1917: 164. Type-species: Carsidaroida heterocephala Crawford, by original 
designation and monotypy. [Synonymised by Crawford, 1920: 355.] 



96 D. HOLLIS 

Description. Integument of head and thorax very sparsely short haired. Head (Figs 7, 8) with disc of vertex 
flat or weakly concave, foveae present as shallow circular depressions or furrows, lateral and hind margins 
of vertex sharply raised, anterior margin sharply defined but weakly incised by median suture, lateral 
ocellae placed posteriorly on vertex, anterolateral tubercles weak or well-developed; antennal sockets 
enlarged, flagellum 2-3-3-0 times longer than head width, first flagellomere more than 12 times longer than 
its median width; genal cones present as a pair of small tubercles on either side of mid ventral line 
immediately anterior to clypeus, weak lateroventral tubercles also present; occiput with a small tubercle on 
each side below eye; ultimate rostral segment long, more than 4 times longer than wide. 

Forewing (Figs 20-24) narrow, about 3 times longer than wide, with subacute apex, costal break absent, 
pterostigma present, r r rs crossvein present in addition to rs-m crossvein, M+Cu stem as long as R stem 
and Cu stem, cuj cell value 1-6 or greater and with radular spinules, apex of claval suture adjacent to apex 
of Cu lb ; hindwing usually with ungrouped costal setae (grouped in T. striata); apical spurs of hind tibia 
arranged 1+2+2. 

Cf Proctiger (Figs 46, 48) unipartite, with well-developed lateral lobes; apex of aedeagus narrow, 
endtube of ductus ejaculatorius slightly swollen (Fig. 50). 

5 proctiger (Figs 81, 82) triangular in profile, usually without posterodorsal lobe (present in T. striata). 

Larva. Undescribed (see Miyatake (1965)). 

Hostplant genera. Sterculia and Heritiera (Sterculiaceae). 

Comments. Like Tenaphalara an additional; r r rs crossvein is present in Tyora and the apical spurs of the 
hind tibia are arranged 1+2+2.1 regard these two characters as autapomorphies for the two genera. Tyora 
differs from Tenaphalara in the raised margins of the vertex, swollen antennal sockets, presence of radular 
spinules in cell cuj, close proximity of the apex of the claval suture to the apex of Cu lb , ungrouped costal 
setae of the hindwing, well-developed lateral lobes of the male proctiger and the absence of a specialised 
posterodorsal lobe on the female proctiger, although a simple conical lobe is present in T. striata. The two 
genera are taken to be the sister-group of the Allocarsidara + Paracarsidara + Mesohomotoma complex. 

Included species 

Further collecting with accurate hostplant data is required before the validity of all the taxa presently 
included in the genus can be adequately assessed. T. ornata and T. striata appear to be good species but it is 
possible that the remainder represents a single species with highly variable forewing pattern. 

Tyora buxtoni Crawford 

(Figs 22, 46, 47) 

Tyora buxtoni Crawford, 1927: 30; 1928: 34; Hodkinson, 1983: 366; Yang & Li, 1985/?: 207, 215. Syntypes, 
2 cf , Samoa [1 cf labelled Type' in BMNH examined]. 

Hostplant. Unknown. 

Recorded distribution. Samoa. 

Material examined 

Further specimens with a similar forewing pattern have been examined from various islands in the 
Solomons group. 

Comments. Although the forewing pattern (Fig. 22) is consistently different from T. congrua, the male and 
female genitalia of buxtoni are morphologically identical to that species. 

Tyora congrua Walker 
(Figs 7, 8, 20, 21, 50, 81) 

Tyora congrua Walker, 1869: 330; Scott, 1882: 471; Aulmann, 1913: 80; Crawford, 1920: 355; Hodkinson, 

1983: 366; Yang & Li, 19856: 206, 215. Holotype cf , Mysol Is [not 'India: Mysore' as in Yang & Li 

(19856)] (BMNH) [examined] . 
Carsidaroida heterocephala Crawford, 1917: 165; 1920: 355. Syntypes cf, ?, Philippines (USNM) 

[examined]. [Synonymised by Crawford, 1920: 355.] 
Nesiope heterocephala var. intermedia Crawford, 1919: 161. Syntypes cf, ? Tanimbar Is, Java [not 

examined]. 



MALVALES-FEEDING PSYLLIDS 97 

Hostplants. Sterculia ureolata, S. lanceolata (Sterculiaceae) (BMNH). 

Recorded distribution. Indonesia (Java, Tanimbar Is, Mysol Is), Philippines. 

Material examined 

Adults from Hong Kong (on Sterculia lanceolata), West Malaysia (Pahang), Indonesia (Java on S. 
ureolata, Sulawesi Utara, Mysol Is), Philippines, Papua New Guinea (Morobe Prov., New Britain on 
Sterculia sp.). 

Comments. Walker's holotype male is very badly damaged and only part of the thorax, one hind leg, the 
abdomen and shreds of wing remain. A microslide preparation of the genitalia, made in 1964, has 
discoloured badly but sufficient remains visible to say that the genitalia are morphologically very similar to 
those of all other specimens examined. The forewing pattern (Figs 20, 21) varies in extent and intensity in 
the material studied, being most dense and extensive in specimens from Sulawesi Utara and least so in the 
Malayan and Hong Kong material. 

Tyora guandongana Yang & Li 

Tyora guandongana Yang & Li, 1985b: 212, 215. Holotype 0\ China: Guandong (BAUIC) [not 
examined]. 

Hostplant. Unknown. 

Comments. From the original figures it appears that this species is not distinct from T. congrua but, as there 
is a dark patch of pattern bordering R+M+Cu stem and R stem (Yang & Li, 19856: fig. 8b) which is not 
present in all material of congrua examined, formal synonymy is not established. 

Tyora ornata (Kirkaldy) 

(Figs 23, 48, 49) 

Nesiope ornata Kirkaldy, 1908: 390; Aulmann, 1913: 81; Laing, 1922: 555; Tuthill, 1964: 357; Miyatake: 

1965: 175; Hodkinson, 1983: 354. Syntypes (? sex), Fiji [not examined]. 
Tyora ornata (Kirkaldy) Crawford, 1920: 355; Caldwell, 1942a: 21; Tuthill, 1951: 273; Yang & Li, 19856: 

206,215. 

Hostplant. Heritiera littoralis (Sterculiaceae). 

Recorded distribution. Ryukyu Is, N. Borneo, S. Marinana Is, Caroline Is, Fiji. 

Material examined 
Adults from Fiji (on H. littoralis), East Malaysia (Sabah). 

Comments. This species is distinguished from T. congrua by the shape of the male paramere and the female 
proctiger and the pattern of the forewing (Figs 23, 49). 

Tyora striata (Crawford) 

(Figs 24, 82) 

Tenaphalara striata Crawford, 1919: 166; Miyatake, 1972: 13; Hodkinson, 1983: 360. Syntypes d\ $, 

Singapore (USNM) [examined]. 
Tyora striata (Crawford) Yang & Li, 19856: 206, 215. 
Tenaphalara sulcata Crawford, 1919: 167, nomen nudum [? in error for striata}. 

Hostplant. Unknown. 

Recorded distribution. Singapore. 

Material examined 

Adults from Singapore and Brunei. 

Comments. The presence of a simple posterodorsal lobe on the female proctiger and the grouped costal 
setae of the hindwing distinguish this species from its congeners. 

TENAPHALARA Kuwayama 

Tenaphalara Kuwayama, 1908: 155; Aulmann, 1913: 75; Crawford, 1919: 163, in part; Mathur, 1975: 165; 
Hodkinson, 1983: 360, in part; 1986: 331, in part; Yang, 1984: 181; White & Hodkinson, 1985: 240, 242; 



98 D. HOLLIS 

Yang & Li, 1985b: 207, 214. Type-species: Tenaphalara acutipennis Kuwayama, by original designation 
and monotypy. 

Description. Integument of head and thorax almost glabrose. Head (Figs 9, 10) with disc of vertex convex, 
foveae present as very shallow circular depressions, hind and lateral margins of vertex rounded, frontal 
margin not sharply defined but deeply incised by median suture, lateral ocellae placed posteriorly on 
vertex, anterolateral tubercles absent; antennal sockets slightly enlarged, antennal flagellum about 2-3 
times longer than head width, 1st flagellomere more than 8 times longer than its median width; 
genal tubercles minute, lateroventral tubercles absent; ultimate rostral segment about 5 times longer than 
wide. 

Forewing (Figs 25, 26) elongate narrow, with subacute apex, about 3-5 times longer than wide, costal 
break absent, pterostigma present, r r rs crossvein present in addition to rs-m crossvein, M+Cu slightly 
longer than R stem and slightly longer than Cu stem, cuj cell value greater than 2-0 and without radular 
spinules, apex of claval suture distant from apex of Cu lb ; hindwing with grouped costal setae; apical spurs 
of hind tibia arranged 1+2+2. 

Cf proctiger (Figs 51, 54) unipartite, with or without lateral lobes; apex of aedeagus narrow, endtube of 
ductus ejaculatorius simple (Figs 53, 56). 

$ proctiger (Fig. 83) with a well-developed dorsal lobe posterior to anal pore. 

Larva. For descriptions of the larval stages of T. acutipennis see Rahman (1932) and Yang (1984). The 5th 
instar larva of T. pseudonervosa is described by Mathur (1973). Wax pore ring arrangement of T. 
camerunus as in Fig. 91. 

Hostplant genera. Bombax, Ceiba (Bombacaceae). 

Comments. Tenaphalara is considered here to be the sister-group of Tyora but has many more derived 
features than the latter. The margins of the vertex are rounded, the antennal sockets are much less swollen, 
the genae bear minute mid ventral tubercles, radular spines are absent from cell cuj of the forewing and the 
apex of the claval suture is distant from the apex of Cu lb , the costal setae of the hindwing are grouped and 
the female proctiger bears a well-developed and specialised posterodorsal lobe. 

The genus has been the most widely misunderstood of the family. The confusion stemming from 
Crawford's (1919) misinterpretation when he included species belonging to other carsidarid genera, e.g. 
malayensis (Crawford) (see p. 101), Juliana (Crawford) (see p. 102) and striata (Crawford) (see p. 97) and 
some non-carsidarid species, e.g. fascipennis (Crawford) (see p. 99) and triozipennis (Crawford) (see p. 
100). Yang & Li (1985b) were aware of this confusion and removed several species from Tenaphalara but 
recombined only fascipennis and striata with more appropriate genera. 

Both included and excluded species are listed below with all the latter recombined. 

Included species 

Tenaphalara acutipennis Kuwayama 

(Figs 25, 51-53) 

Tenaphalara acutipennis Kuwayama, 1908: 156; Crawford, 1911: 491; 1919: 164 [possible misidentifica- 

tion]; Aulmann, 1913: 75; Ramakrishna Ayyar, 1924: 623; Boselli, 1930: 188; Kuwayama, 1931: 125; 

Mathur, 1935: 63; 1975: 166; Beeson, 1941: 595; Heslop-Harrison, 1951: 26; Sohi & Sandhu, 1978: 237; 

Hodkinson, 1983: 360; 1986: 331; Yang, 1984: 181; Yang & Li, 1985b: 206, 214. Syntypes cf, $, Japan, 

? Taiwan [not examined]. 
Tenaphalara elongata Crawford, 1912: 432; 1919: 164; Rahman, 1932: 367; Yang & Li, 1985b: 206, 214. 

Syntypes cf , ?, India: on 'Silk cotton' [not examined]. [Synonymised by Crawford, 1919: 164.] 
Ctenophalara elongata Bhatia & Shaffi, 1933: 545 etseq. [Misspelling.]. 

Hostplants. Bombax ceiba (Bombacaceae). Crawford's (1919) record on Sterculia foetida and BMNH 
specimens on Hibiscus esculentus are regarded as doubtful host records. 

Recorded distribution. India, Taiwan, Japan, Philippines. 

Material examined 

Adults and larvae from Pakistan (on Bombax ceiba), India, Bangladesh (on Bombax ceiba and Hibiscus 

esculentus) . 

Comments. Most authors have accepted Crawford's synonymy of elongata with acutipennis but Yang & Li 
(1985b), basing their judgement on a combination of Kuwayama's (1908) and Mathur's (1975) figures, 



MALVALES-FEEDING PSYLLIDS 99 

regarded acutipennis and elongata as separate species. Evidence from the BMNH collections suggests that 
only one species is involved. 

Tenaphalara camerunus (Aulmann) comb. n. 

(Figs 9, 10,26,54-56,83) 

Carsidara camerunus Aulmann, 1912/?: 21; 1913: 79. Holotype cf, Cameroon (MNHU). 

Hostplants. Ceiba pentandra, Bombax buonopozense and B. sessile (Bombacaceae). 

Material examined 

Adults and larvae from Senegal, Sierra Leone, Ghana, Nigeria, Cameroon, Zaire and Zanzibar. 

Comments. Although the holotype has not been examined there are several samples of this species in 
BMNH collections; a study of these and the original description has shown that camerunus should be 
placed in Tenaphalara. The species is similar to acutipennis but differs in that the apical segment of the 
aedeagus is relatively shorter (Fig. 56), the male proctiger has weakly developed lateral lobes (Fig. 54), the 
paramere is of a different shape (Fig. 55) and the female subgenital plate is trilobed apically (Fig. 83). 

Tenaphalara gossampini Yang & Li 

Tenaphalara gossampini Yang & Li, 19856: 208. Holotype cf, China: on Bombax ceiba [as 'Gossampinus 
malabarica\ (BAUIC) [not examined]. 

Comments. From the original figures (Yang & Li, 1985/?) of this and the following three species, it is likely 
that all four are synonyms of T. acutipennis but formal synonymy is not established without reference to 
type material. No larvae were mentioned in the original data for all these species and it is improbable that 
Aphanamixis grandifolia (Meliaceae), Mangifera indica (Anacardiaceae) or Dimocarpus longana (Sapin- 
daceae) are true hostplants. 

Tenaphalara aphanmixis Yang & Li 

Tenaphalara aphanmixis Yang & Li, 19856: 209. Holotype d\ China: on Aphanamixis grandifolia 
(BAUIC) [not examined]. 

Tenaphalara dimocarpi Yang & Li 

Tenaphalara dimocarpi Yang & Li, 19856: 211. Holotype J, China: on Dimocarpus longana (BAUIC) 
[not examined]. 

Tenaphalara mangiferae Yang & Li 

Tenaphalara mangifera Yang & Li, 19856: 210. Holotype 0\ China: on Mangifera indica (BAUIC) [not 
examined]. 

Tenaphalara pseudonervosa (Mathur) comb. n. 

Nesiope pseudonervosa Mathur, 1973: 71; Hodkinson, 1986: 319. Holotype d\ India: on Bombax ceiba 

(FRI) [not examined]. 
Ty or a pseudonervosa (Mathur) Yang & Li, 19856: 207, 215. 

Host plant. Bombax ceiba (Bombacaceae). 

Comments. This new combination is made on the basis of the original description ; the type material has not 
been examined. Mathur's figures show a convex vertex, absence of radular spinules from cell ciij and the 
characteristically specialised posterodorsal lobe of the female proctiger. 

Excluded species 

Strogylocephala fascipennis Crawford 

Strogylocephalafascipennis Crawford, 1917: 167; Yang & Li, 19856: 207, 215. Syntypes G\ 2 , Philippines 

(USNM) [examined, all damaged]. 
Tenaphalara fascipennis (Crawford) Crawford, 1919: 165; Miyatake, 1971: 68. 



100 D. HOLLIS 

Comments. Crawford (1919) placed this Philippines species in Tenaphalara and synonymised Strogy- 
locephala with Tenaphalara. This synonymy was accepted by some subsequent authors (Yu, 1957; 
Miyayake, 1971) but Yang & Li (19856) disputed this and regarded Strogylocephala as a valid genus. Yu 
(1957) described Tenaphalara confluens from Taiwan, on Palaquium formosanum (Sapotaceae), and 
suggested it was closely related to fascipennis . Yang & Li (1985/?) placed confluens also in Strogylocephala. 
Chung Tu Yang (1984) redescribed confluens and placed it in a new genus Synaphalara, in the subfamily 
Carsidarinae. 

After examining Crawford's type material of fascipennis and recently collected specimens of the same 
species from Sulawesi Utara, on Palaquium sp. I agree with Yang & Li (19856) that Strogylocephala is a 
distinct genus. It is quite unrelated to the Carsidaridae and I tentatively place it in the Calophyidae. 
Furthermore I consider Synaphalara to be a junior synonym of Strogylocephala (syn. n.) and it is likely that 
confluens and fascipennis are synonyms. 

Placing Strogylocephala in the Calophyidae conflicts with published data on the larva of the type-species 
which was clearly described as a triozid larva by Chung Tu Yang (1984: 186). However, when the Sulawesi 
specimens of fascipennis were collected, larvae, which agree generally with Yang's description, and a single 
adult triozid were taken at the same time. This triozid is an undescribed Trioza species belonging to the 
Sapotaceae/Ebenaceae-feeding obsoleta-group (Hollis, 1984). It is therefore possible that the larvae 
described by Yang belong to this triozid species and are not associated with the Strogylocephala species. 

Strogylocephala confluens (Yu) 

Tenaphalara confluens Yu, 1956: 46. Holotype cf , Taiwan [not examined]. 
Synaphalara confluens (Yu) Chung Tu Yang, 1984: 185. 
Strogylocephala confluens (Yu) Yang & Li, 19856: 207, 216. 

Comments. See under S. fascipennis. 

Allocarsidara Juliana (Crawford) comb, n.; see below (p. 102) 

AHocarsidara malayensis (Crawford) comb, n.; see below (p. 101) 

Tenaphalara sulcata Crawford, 1919: 167, nomen nudum; see above (p. 97) 

Trioza triozipennis (Crawford) comb. n. 

Tenaphalara triozipennis Crawford, 1919: 167. Holotype ?sex [abdomen missing], Singapore (USNM) 

[examined] . 
? Tenaphalara triozipennis Crawford; Yang & Li, 19856: 207, 216. 

Comments. The holotype is clearly a triozid, probably related to T. umalii (Miyatake) (see below), and is 
provisionally placed in Trioza. 

Trioza umalii (Miyatake) comb. n. 

Tenaphalara umalii Miyatake, 1972: 11. Holotype $, Philippines (OMNH) [not examined]. 
? Tenaphalara umalii Miyatake; Yang & Li, 19856: 207, 216. 

Comments. From Miyatake's description there can be little doubt that this species is a triozid and is 
probably related to triozipennis (Crawford). The above new combination is therefore proposed. 

ALLOCARSIDARA gen. n. 

[Tenaphalara Crawford, 1919: 163, in part; Hodkinson, 1983: 360, in part; 1986: 331, in part; Braza & 

Calilung, 1981: 354. Misinterpretations.] 
Type-species: Tenaphalara malayensis Crawford. 

Description. Integument of head and thorax almost glabrous. Head (Figs 11, 12) with disc of vertex flat or 
convex, foveae weak, lateral and hind margins of vertex rounded, anterior margin not clearly defined and 
weakly incised by median suture, lateral ocelli placed medially on vertex, anterolateral tubercles 
well-developed; antennal sockets enlarged, flagellum 3-26-4-60 times longer than head width, first 
flagellomere more than 10 times longer than wide; genal cones present as a pair of very weakly developed 



MALVALES-FEEDING PSYLLIDS 



101 



humps on either side of mid ventral line anterior to clypeus, genae also with an outer pair of lateroventral 
tubercles, occiput with a small tubercle on each side below eye; ultimate rostral segment more than four 
times longer than wide. 

Forewing (Figs 29-34) narrow elongate , at least 2-8 times longer than wide and with an acute apex, costal 
break absent, pterostigma present or absent, r r rs crossvein absent, M+Cu stem as long as R stem and 
more than 1-7 times longer than Cu stem, cuj cell value 1 -76-2-70 and without radular spinules, apex of 
claval suture distant from apex of Cu lh ; hindwing with grouped costal setae; apical spurs of hind tibia 
arranged 1+2+1. 

Cf proctiger (Figs 57, 60, 63, 66, 69, 72) unipartite, with well-developed lateral lobes that are sometimes 
subdivided posteroapically, each inner lateral surface with a patch of differentiated spinules; apical 
segment of aedeagus straight or flexed, aedeagus narrow apically with endtube of ductus ejaculatorius 
contained within apical lobes (Figs 59, 62, 65, 68, 71, 74). 

$ proctiger (Figs 84-87) with or without a posterodorsal lobe. 

Larva. As in Fig. 92. 

Hostplant genus. Durio (Bombacaceae). 

Comments. The sister-group relationship of Allocarsidara needs to be resolved. The genus is placed with 
Paracarsidara and Mesohomotoma on the structure of the male proctiger but it differs from these genera in 
having rounded lateral and posterior margins of the vertex, well-developed anterolateral tubercles on the 
vertex, a simple endtube of the ductus ejaculatorius that is enclosed within the apical lobes of the aedeagus. 
and a patch of differentiated spinules on each inner lateral surface of the male proctiger. Some species have 
lost the pterostigma of the forewing as in Mesohomotoma. 

Yang & Li (1985b) expressed doubt that Tenaphalara malayensis Crawford and T. Juliana Crawford 
belonged in Tenaphalara and this suggestion is confirmed here. Crawford (1919) misinterpreted the genus 
Tenaphalara (see above, p. 98) and his two species malayensis and Juliana are transferred to Allocarsi- 
dara. A further four new species are described below. 

Relationships among these species are not at all clear. A variety of characters appear in the group, e.g. 
the absence of a pterostigma, an apically bifid male paramere, a flexed distal aedeagal segment, the male 
proctiger with subdivided lateral lobes, and the female proctiger with a posterodorsal lobe. However, the 
distribution of these characters within the genus (Table 1) gives little indication of sister-group rela- 
tionships. The six species included in Allocarsidara may be separated from one another by using the 
character-matrix in Table 1; three are known to develop on Durio zibethinus. 

Table 1 Allocarsidara spp. , character-matrix. Primitive condition given in parenthesis. 







Posterodorsal 












lobe of ? 




apical segment 


lateral lobes 




pterostigma 


proctiger 


Cf paramere 


of aedeagus 


of cf proctiger 




absent 


present 


truncate 


flexed 


divided 


Character 


(present) 


(absent) 


(conical) 


(straight) 


(entire) 


Species 












malayensis 


(-) 


+ 


+ 


(") 


(-) 


iriana 


(") 


? 


+ 


(") 


(-) 


incognita 


(") 


+ 


(-) 


+ 


+ 


bakeri 


+ 


? 


(") 


+ 


(-) 


Juliana 


+ 


I 1 ) 


+ 


+ 


(-) 


elongata 


+ 


+ 


(") 


(") 


(-) 



Included species 

Allocarsidara malayensis (Crawford) comb. n. 

(Figs 29, 57-59, 86) 

Tenaphalara malayensis Crawford, 1919: 165; 1928: 425; Laing, 1930: 39; Hodkinson, 1983: 360; 1986: 332. 

Syntypes, cf , 5, Malaya (Penang) and Singapore (USNM) [examined]. 
? Tenaphalara malayensis Crawford; Yang & Li, 1985b: 207, 216. 

Hostplant. Durio zibethinus (Bombacaceae). 



102 D. HOLLIS 

Recorded distribution. Malaya, Singapore, Sumatra, Java. Braza & Calilung (1981: 355) record this 
species from the Philippines but their material represents a different species (see below). 

Material examined 

Adults and larvae from Thailand, Malaya and Java. 



Allocarsidar a Juliana (Crawford) comb. n. 

(Figs 32, 66-68, 84) 

Tenaphalara Juliana Crawford, 1919: 166; 1920: 355; Hodkinson, 1983: 360. Holotype, ? sex, 'Borneo' 

(BPBM) [examined]. 
? Tenaphalara Juliana Crawford; Yang & Li, 19856: 207, 216. 

Larva and hostplant. Unknown. 

Recorded distribution. Sabah. 

Material examined 

Holotype (? sex), East Malaysia: Sabah ('Sandakan, Borneo, Baker'); 1 o\ 1 $, same data as holotype 
(USNM); 2 $> , Sarawak, offshore gas platform (BMNH). 

Comments. This species was described from a single specimen of unknown sex (the posterior half of the 
abdomen was missing) from Sabah, and further recorded (Crawford, 1920) from a single female from an 
unstated locality but presumably in Borneo as the paper in which the specimen is recorded deals with 
Bornean psyllids. Apart from the holotype (BPBM), there are three specimens bearing the type data, 
under the name 'Tenaphalara Juliana, in USNM; two complete males and a complete female. The two 
males are not conspecific with each other and neither can be positively identified as the same species as the 
damaged holotype of Juliana. The complete female agrees well with the holotype and I assume that it is the 
specimen Crawford recorded as Tenaphalara Juliana in 1920. As well as the locality label this specimen also 
bears a handwritten label '11734', which is almost certainly an original Baker collection label. One of the 
two males also bears a similar label and, therefore, I regard it as having been collected by Baker at the same 
time as the female. I consider this specimen to be the male of Allocarsidara Juliana and figure it below (Figs 
32, 66-68). The other male in USNM is described below as A. bakeri sp. n. (p. 103). 

Allocarsidara Juliana differs from malayensis in lacking a pterostigma in the forewing (Fig. 32), lacking a 
posterodorsal lobe on the female proctiger (Fig. 84) and having a flexed apical aedeagal segment in the 
male (Fig. 68). It is apparently closest to bakeri (see below) but differs from that species in the shape of the 
paramere (Figs 67, 70). 



Allocarsidara incognita sp. n. 

(Figs 30, 72-74, 85, 92) 

[Tenaphalara malayensis Crawford; Braza & Calilung, 1981: 355. Misidentification.] 

Description (only slide-mounted and spirit-stored material available for study). Antennal flagellum 
3-83-4-30 times longer than head width, 1st flagellomere more than 13 times longer than wide; ultimate 
rostral segment 1-52-2-17 times longer than flagellomeres 7+8 together. 

Forewing (Fig. 30) 2-99-3-18 times longer than wide, pterostigma present, Cu stem 3 times longer than 
Cu lb , cuj cell value 1-83-2-62. 

Cf genitalia as in Figs 72-74; lateral lobes of proctiger each with a supplementary apicoposterior lobe; 
paramere narrow apically and with an inwardly directed spine, apical segment of aedeagus flexed. 

5 terminalia as in Fig. 85, proctiger with a well-developed posterodorsal lobe. 

Measurements (7 d\ 7 $). Maximum width of head, cf 0-67-0-71, $ 0-71-0-74; length of antennal 
flagellum, cf 2-67-2-90, $ 2-74-3-06; length of ultimate rostral segment, cf 0-26-0-30, $ 0-30-0-31; 
length of forewing, cf 3-32-3-53, $ 3-65-3-89; length of hind tibia, cf 0-68-0-76, $ 0-71-0-74. 

Larva. Abdominal wax pore rings as in Fig. 92. 

Hostplant. Adults and larvae collected from Durio zibethinus (Bombacaceae). 



MALVALES-FEEDING PSYLLIDS 103 

Holotype d\ Indonesia: Sulawesi Utara, Dumoga Bone NP, Bungalows light trap, 21. xi. 1985 (RESL 
Project Wallace) (BMNH); slide-mounted. 

Paratypes. Indonesia: 1 J , same locality data as holotype, Base Camp light trap, 26.x. 1985. Philippines: 
1 J, DavaoExp. Stn, ground level trap, xi. 1962 (M. R. Gavarra);2 cf, 2 $, larvae, DavaoCity, 2.x. 1975, 
Durio zibethinus (M. Gavarra). East Malaysia: 5 cf, 4 J, Sarawak, vi. 1963, Maize (C. R. Wallace) 
(BMNH). 

Comments. This species is very similar to malayensis but can be recognised by the structure of the male 
genitalia; females of the two species are very difficult to separate from one another. 

Allocarsidara iriana sp. n. 

(Figs 11-13, 31, 63-65) 

Description. Body mid brown dorsally and laterally, ochraceous ventrally; head black but with a median 
longitudinal pale stripe on vertex, apices of anterolateral tubercles ochraceous; antenna with scape and 
pedicel ochraceous, flagellum black; forewing hyaline; femora ochraceous, brown anterodorsally, fore and 
mid tibiae and tarsi dark brown, hind tibia and tarsus ochraceous. 

Antennal flagellum relatively short, 3-26 times longer than head width, first fiagellomere 10 times longer 
than its greatest width, seventh fiagellomere with a very long subapical seta (Fig. 13); ultimate rostral 
segment 1-46 times longer than flagellomeres 7+8 together. 

Forewing (Fig. 31) 2-86 times longer than wide, pterostigma present, M+Cu stem just over twice as long 
as Cu stem, Cu stem twice as long as Cu lb , cu } value 2-13. 

Cf genitalia as in Figs 63-65; lateral lobes of proctiger short, not subdivided posteroapically; paramere 
broad apically, with an anteriorly directed spine and an apicoposterior tubercle; apical segment of 
aedeagus straight. 

9 unknown. 

Measurements (1 cf). Maximum width of head, 0-58; length of antennal flagellum, 1-88; length of 
ultimate rostral segment, 0-23; length of forewing, 3-00; length of hind tibia, 0-76. 

Larva and hosplant. Unknown. 

Holotype cf , New Guinea: NW, Nabire, S. Goelvink Bay, 7.ix.l962, light trap in jungle (H. Holtman) 
(BPBM); slide-mounted. 

Comments. The forewing of this species has a pterostigma, as in malayensis and incognita, but it may be 
recognised by the shape of the male paramere and aedeagus, the relatively short antennal flagellum and the 
long subapical seta on fiagellomere seven. 

Allocarsidara baker isp. n. 

(Figs 33, 69-71) 

Description. Body light brown, head darker; antenna with scape, pedicel and first fiagellomere dark 
brown, rest of flagellum lighter; forewing hyaline; legs ochraceous, fore and mid femora with dark 
anterodorsal stripes, mid tibia and tarsus darker. 

Antennal flagellum 4-66 times longer than head width, first fiagellomere 12 times longer than its greatest 
width. 

Forewing (Fig. 33) 3-45 times longer than wide, pterostigma absent, Cu stem 3-5 times longer than Cu lh , 
cuj cell value 2-7. 

Cf genitalia as in Figs 69-71; lateral lobes of proctiger long but not subdivided posteroapically; apex of 
paramere narrow and with an inwardly directed curved spine; apical segment of aedeagus flexed. 

$ unknown. 

Measurements (1 cf). Maximum width of head, 0-59; length of antennal flagellum, 2-77; length of 
ultimate rostral segment, 0-24; length of forewing, 3-12; length of hind tibia, 0-57. 

Larva and hostplant. Unknown. 

Holotype cf , East Malaysia: Sabah; k Sandakan, Borneo, Baker (USNM); slide-mounted. 

Comments. This species is very similar to Juliana but differs in that the lateral lobes of the male proctiger are 
longer (Figs 66, 69), the paramere is narrow apically (Figs 67, 70), the apex of the aedeagus is of a different 
shape (Figs 68, 71) and the forewing is relatively narrower (3-10 times longer than wide in Juliana). 



104 D. HOLLIS 

Allocarsidara elongata sp. n. 

(Figs 34, 60-62, 87) 

Description (only slide-mounted and spirit-preserved material available for study). Antennal flagellum 
3-98-4-60 times longer than head width, first flagellomere 16 times longer than its greatest width; ultimate 
rostral segment relatively long, 2-54-2-89 times longer than flagellomeres 7+8 together. 

Forewing (Fig. 34) 3-21-3-35 times longer than wide, pterostigma absent, Cu stem about 4 times longer 
than Cu lb , cu 1 cell very small. 

d" genitalia as in Figs 60-62; lateral lobes of proctiger undivided posteroapically; apex of paramere 
narrow, with a large and inwardly directed apical tubercle; apical segment of aedeagus straight. 

5 terminalia as in Fig. 87, proctiger with a well-developed posterodorsal lobe. 

Measurements (6 cf , 7 $). Maximum width of head, cf 0-64-0-73, $ 0-69-0-75; length of antennal 
flagellum, cf 2-83-3-00, $ 3-11-3-26; length of ultimate rostral segment, cf and $ 0-31-0-33; length of 
forewing, Cf 3-47-3-59, $ 3-81-4-14; length of hind tibia, cf 0- 69-0-73, $ 0-71-0-74. 

Hostplant. Adults and larvae collected from Durio zibethinus (Bombacaceae). 

Holotype cf, West Malaysia: Malaya, Alor Star, Jitra, 3.xii.l985, Durio zibethinus (N. K. Ho) 
(BMNH); slide-mounted. 

Paratypes. 7 cf , 9 $ , same data and depository as holotype; slide- and dry-mounted. 

Comments. This species resembles Juliana and bakeri in lacking a pterostigma in the forewing; it differs 
from both in having a straight apical aedeagal segment, and from Juliana in having an apically narrow 
paramere and a well-developed posterodorsal lobe on the female proctiger. 



PARACARSIDARA Heslop-Harrison 

[Carsidara Walker; Crawford, 1911: 484; 1914: 57; Tuthill & Taylor, 1950: 53; Brues et ai, 1954: 156; 
Morgan, 1984: 19, fig. 15. Misinterpretations.] 

Paracarsidara Heslop-Harrison, 1960: 244; Hodkinson & White, 1981: 505; Brown, 1985; 246. Type- 
species: Carsidara dugesii Low, by original designation. 

Description. Integument of head and thorax almost glabrose. Head (Figs 14, 15) with disc of vertex 
concave, foveae present as deep oblique grooves, lateral margins of vertex sharply raised, posterior margin 
angular, anterior margin sharply defined and deeply incised by median suture, lateral ocellae placed 
posteriorly on vertex, anterolateral tubercles very weak or absent; antennal sockets swollen, flagellum 
2-5-3-5 times longer than head width, 1st flagellomere more than 10 times longer than its greatest width; 
genae with a pair of small mid ventral tubercles anterior to clypeus, each gena with a larger lateroventral 
tubercle; occiput, on each side below, eye with a small tubercle or swelling; ultimate rostral segment more 
than 6 times longer than wide. 

Forewing (Fig. 35) narrow-elongate, about 2-8 times longer than wide and with a subacute apex, costal 
break absent, pterostigma present, r r rs crossvein absent, M+Cu stem almost as long as R stem and Cu 
stem, cuj cell value greater than 2-7 and without radular spinules, apex of claval suture distant from apex of 
Cu lb , costal setae of hindwing grouped; apical spurs of hind tibia arranged 1+3 + 1. 

Cf proctiger (Fig. 75) unipartite, expanded basally and with 2 small apicolateral lobes; aedeagus narrow 
apically, sometimes lobed basoventrally, endtube of djuctus ejaculatorius enlarged and with lateral and 
apical hooks (Fig. 76). 

$ terminalia (Fig. 88) conical, proctiger more or less 'stepped' posteriorly, apical part narrow and 
bearing short, thick, spiniform setae and sometimes an apical hook. 

Larva. Caudal plate as in Fig. 93. 

Hostplant genera. Ceiba (Bombacaceae), Malva and Wissadula (Malvaceae). 

Comments. On the basis of the narrow-elongate forewing, absence of r r rs crossvein, long M+Cu stem, 
presence of lateral lobes on the male proctiger and narrow apex of the aedeagus, Paracarsidara is grouped 
with Allocarsidara and Mesohomotoma. The modified endtube of Paracarsidara is very similar to that of 
Mesohomotoma and is used as a synapomorphy for these two genera. Brown (1985) also related 
Paracarsidara to Mesohomotoma, even doubting their separate identities, but I regard the absence of a 
pterostigma and the bipartite male proctiger that bears a median posterior lobe in addition to the lateral 
lobes sufficient to recognise the latter genus. 



MALVALES-FEEDING PSYLLIDS 105 

Included species 
Paracarsidara dugesii (Low) 

(Figs 75, 76, 88, 93) 

Carsidara dugesii Low, 1886: 160; Crawford, 1911: 486; 1914: 58; Aulmann, 1913: 79; Laing, 1923: 698; 

Caldwell, 1941: 421, 1944: 58; Tuthill, 1950: 53; Silva etal. , 1968: 200 [?misidentification]. Syntypes 0\ 

$ , Mexico: 'planta Malvacea' (NM) [not examined]. 
Paracarsidara dugesii (Low) Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 506; Brown, 1985: 

250. 
Carsidara concolor Crawford, 1911: 484; 1914: 58; Caldwell, 1942/?: 28; Caldwell & Martorell, 1951: 604. 

Syntypes c? , $, Cuba (USNM). [Synonymised by Heslop-Harrison, 1960: 246.] 
Paracarsidara concolor (Crawford) Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 505. 
[Carsidara gigantea Crawford; Laing, 1923: 698. Misidentification.] 

Hostplants. Malva sp., Wissadula perplocifolia (Malvaceae). Silva et al. (1968) record this species from 
Brazil, on Bombax cyathophorum (Bombacaceae), but this is likely to be a misidentification. 

Recorded distribution. Mexico, Guatemala, Cuba, Puerto Rico, Dominica and Jamaica; [?Brazil]. 

Material examined 

Adults and larvae from Mexico and Jamaica (on Wissadula perplocifolia). 

Paracarsidara gigantea (Crawford) 
(Figs 14, 15,35) 

Carsidara gigantea Crawford, 1911: 486; 1914: 57; Ferris, 1928: 244; Caldwell, 1941: 421. Syntypes $, 

Nicaragua (USNM). 
Paracarsidara gigantea (Crawford) Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 506; Brown, 

1985:253. 

Hostplants. Ceiba sp., C. burchellii (BMNH). 

Recorded distribution. Mexico, Guatemala, Belize, Nicaragua, St Kitts, Cuba, Colombia, Brazil. 

Material examined 

Adults and larvae from Mexico, Panama and Brazil (on Ceiba burchellii). 

Comments. Tuthill (1950) synonymised this species with dugesii but Brown (1985) considered it to be 
distinct after examining relevant type material. 

Paracarsidara rostrata (Crawford) 

Carsidara rostrata Crawford, 1911: 486; 1914: 58; Tuthill, 1950: 53. Syntypes $, Nicaragua (USNM). 
Paracarsidara rostrata (Crawford) Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 506; Brown, 
1985:256. 

Hostplant. Unknown. 

Recorded distribution. Nicaragua, Panama. 

Comments. Crawford (1914) regarded this species as a synonym of dugesii and this was followed by Tuthill 
(1950). Heslop-Harrison raised rostrata from synonymy and Brown (1985) confirmed this after examining 
type material. 

Nomen dubium 

Carsidara mexicana Crawford, 1911: 487; 1914: 58 (as a possible synonym of dugesii). Holotype d\ Mexico 

[specimen missing from pin, USNM]. 
Paracarsidara mexicana (Crawford); Heslop-Harrison, 1960: 246; Hodkinson & White, 1981: 248 (as a 

nomen dubium). 

Undescribed species 

Brown (1985) recorded two undescribed species from Panama and there are some poorly preserved 
specimens of a further undescribed species from Trinidad on Malachra alceifolia (Malvaceae) in BMNH. 



106 D. HOLLIS 

MESOHOMOTOMA Kuwayama 

Mesohomotoma Kuwayama, 1908: 180; Aulmann, 1913: 36; Crawford, 1920: 356; 1925a: 32; Tuthill & 
Taylor, 1955: 249; Miyatake, 1971: 58; Mathur, 1975: 144; Braza & Calilung, 1981: 356; Hodkinson, 
1983: 352; 1986: 317; Yang, 1984: 178; White & Hodkinson, 1985: 242. Type-species: Mesohomotoma 
camphorae Kuwayama, by original designation and monotypy. 

Udamostigma Enderlein, 1910: 138. Type-species: Tyora hibisci Froggatt, by original designation and 
monotypy. [Synonymised by Crawford, 1920: 356.] 

[Tyora Walker; Crawford, 1919: 159. Misinterpretation.] 

[ Ty orfija Walker; Uichanco, 1921: 279. Misinterpretation; misspelling.] 

Description. Integument of head and thorax almost glabrous. Head (Figs 16, 17) with disc of vertex deeply 
divided by median suture, foveae present as deep oblique grooves, lateral and hind margins of vertex 
obliquely raised, anterior margin poorly defined but deeply incised by median suture, anterolateral 
tubercles well-defined, lateral ocellae posteriorly placed; antennal sockets enlarged, flagellum 2-4-3-0 
times longer than head width, 1st flagellomere at least 9 times longer than its median width; genae with a 
small tubercle on either side of mid ventral suture immediately anterior to clypeus, lateral tubercles absent; 
occiput with a small tubercle on each side below eye; ultimate rostral segment at least 5 times longer than 
wide. 

Forewing (Figs 36-39) narrow, elongate, 2-5-3-0 times longer than wide, with subacute apex; costal 
break, pterostigma and r r rs absent, M+Cu stem short, about two-fifths as long as R stem and half as long 
as Cu stem, cuj cell value almost 2-0 and without radular spinules, Cu la strongly arched towards M stem, 
apex of claval suture distant from apex of Cu lb ; costal setae of hindwing grouped; apical spurs of hind tibia 
arranged 1+3 + 1. 

Cf proctiger (Fig. 77) bipartite, a strong anvil-shaped median posterior lobe present in addition to 
well-developed lateral lobes; aedeagus (Fig. 78) narrow apically, apex of ductus ejaculatorius prominent, 
strongly produced from aedeagal apex and expanded apically. 

9 proctiger (Fig. 89), in profile, strongly stepped posteriorly, apical part narrow elongate and bearing 
short and thickened setae, apex weakly barbed. 

Larva. Caudal plate as in Fig. 94. Larval stages of M. camphorae described by Yang (1984). 

Hostplant genera. Hibiscus , Thespesia, Abutilon, Urena (Malvaceae); Cola, Theobroma, Sterculia, 
Heritiera (Sterculiaceae). 

Comments. Mesohomotoma differs from all other carsidarids in having a bipartite male proctiger that bears 
a median posterior lobe in addition to the lateral lobes. Like Carsidara the M+Cu stem is short but this is 
probably a primitive feature; the pterostigma is absent, resembling some species of Allocarsidara but this is 
likely to be a parallel development in the two genera; the form of the apical segment of the aedeagus and 
particularly the endtube of the ductus ejaculatorius is similar to Paracarsidara and this is considered as a 
synapomorphy for the two genera. 

Included species 

A great deal of confusion exists over the identity and validity of the six species currently recognised in this 
genus. I have not examined original type-material but have seen a large number of specimens from most of 
the areas from where these species were described. There is considerable variation in size, body coloration 
and forewing pattern and vein proportions (Figs 36-39) between, but not within, samples, although this 
variation does not correlate well with hostplant data or geographical distribution. Male and female 
genitalia and larval morphology are, however, remarkably consistent throughout the geographical and 
hostplant ranges of the material. 

The most commonly encountered species is M. hibisci and its hostplant, Hibiscus tiliaceus, is common in 
strandline and coastal vegetation throughout the Pacific and Indian Ocean regions from Polynesia to the 
East and South African coast. M. camphorae and M. lineaticollis , both originally described from Taiwan, 
have been subsequently recorded on Hibiscus tiliaceus and other malvaceous hosts such as Urena lobata 
and Abutilon indicum, in South East Asia. M. lutheri, described from Sri Lanka, is recorded from Hibiscus 
tiliaceus and Urena lobata in India, Sri Lanka, Reunion and Madagascar. M. africana was described from 
Natal without host data but there are Mesohomotoma samples in BMNH collected from Hibiscus in eastern 
Tanzania, Kenya, Uganda and Zaire; M. tessmanni is widespread in Central and West Africa on Cola, 
Theobroma (cocoa) and other sterculiaceous hosts. 

I suspect that camphorae, lineaticollis, lutheri and africana are synonyms of hibisci. Furthermore, there 
appear no consistent morphological characters, either in the adult or larva, that will separate tessmanni 



MALVALES-FEEDING PSYLLIDS 107 

from hibisci but the sterculiaceous hostplant preferences of the West and Central African populations 
suggest some genetic isolation from the Malvaceae-feeding populations. Hibiscus tiliaceus does occur 
around the coastal areas of West Africa but it is not known if these bushes support populations of 
Mesohomotoma. Further collecting here would be useful. Experimental work involving hostplant transfers 
and crossbreeding techniques is required to clarify the problem. 

Mesohomotoma hibisci (Froggatt) 
(Figs 36, 37, 77, 78, 89, 94) 

Tyora hibisci Froggatt, 1901: 287. Syntypes cf , $, Australia: Queensland, on Hibiscus tiliaceus (ANIC) 

[not examined]. 
Udamostigma hibisci (Froggatt) Enderlein, 1910: 138; Aulmann, 1913: 81. 
Mesohomotoma hibisci (Froggatt) Crawford, 1920: 356; 1925a: 34; 1927: 30; Klyver, 1933: 25; Caldwell, 

1942: 21; Tuthill, 1943: 71; 1951: 273; 1964: 355; Tuthill & Taylor, 1955: 250; Hodkinson, 1983: 353; 

1986: 317; Morgan, 1984: 29. 

Hostplants. Hibiscus tiliaceus, H. rosasinensis, H. boryanus (Malvaceae). 

Recorded distribution. Polynesia (Society, Austral and Rapa Is), Melanesia (Fiji and Solomon Is, New 
Caledonia), Micronesia (Marianas and Caroline Is), Australia (Queensland). 

Material examined 

Numerous adults and larvae from: Polynesia (Cook, Society, Tonga, Samoa and Ellice Is), Melanesia 
(Fiji, Loyalty, New Caledonia, Solomons, Bismark Is and Papua New Guinea), Micronesia (Caroline, 
Marianas and Gilbert Is), Australia (N.S.W.), Indonesia (Sulawesi), Philippines, Hong Kong, India. 
Chagos Arch., Seychelles, Reunion, Mauritius, Madagascar, Zaire, Uganda, Kenya, Tanzania, Zimbabwe, 
South Africa (Natal). 

Mesohomotoma africana Pettey 

Mesohomotoma africana Pettey, 1924: 29; 1925: 138; Capener, 1970: 199. Syntypes cf, $, South Africa 
(Natal) (NCI; BMNH) [1 £ examined]. 

Hostplant. Hibiscus sp. (Malvaceae). 

Recorded distribution. South Africa (Natal). 

Mesohomotoma camphor ae Kuwayama 

Mesohomotoma camphorae Kuwayama, 1908: 181; Aulmann, 1913: 36; Crawford, 1925a: 33; 1927: 31; 
1928: 34; Kuwayama, 1931: 123; Takahashi, 1936: 292; Sasaki, 1954: 32; Miyatake, 1964: 123; 1965: 174; 
1971: 58; Hodkinson, 1983: 352; 1986: 317; Yang, 1984: 178. Syntypes d\ $, Japan (Osagawara Is) and 
Taiwan, on 'Kampferbaume' [not examined]. 

Hostplants. Hibiscus tiliaceus, H. mutabilis, Abutilon indicum, Thespesia populnea, Urena lobata 
(Malvaceae); Cinnamomum camphora (Lauraceae) but Sasaki (1954) regards the camphor records as 
erroneous. 

Recorded distribution. Samoa, Fiji, Philippines, Japan (Osagawara and Ryukyu Is), Taiwan. 

Mesohomotoma lineaticollis Enderlein 

Mesohomotoma lineaticollis Enderlein, 1914: 232; Crawford, 1925a: 35; Boselli, 1930: 188; Kuwayama, 
1931: 123; Braza & Calilung, 1981: 356; Hodkinson, 1983: 353. Holotype cf , 'Formosa' [not examined]. 

Hostplant. Urena lobata (Malvaceae). Also recorded (probably in error) on Cordia dichotoma (Boragi- 
naceae). 

Recorded distribution. Taiwan, Philippines. 

Mesohomotoma lutheri (Enderlein) 

Udamostigma lutheri Enderlein, 1918: 484. Syntypes cf , §, Sri Lanka [not examined]. 
Tyora indica Crawford, 1919: 159. Syntypes cf , $, Amboina and India [not examined]. [Synonymised by 
Crawford, 1925a: 34.] 



108 D. HOLLIS 

Tyorfija indica Crawford; Uichanco, 1921: 279. [Misspelling.] 

Mesohomotoma lutheri (Enderlein) Crawford, 1925a: 34; 1927: 31; Mathur, 1975: 146; Hodkinson, 1983: 

353; 1986: 317. 
Mesohomfajtoma lutheri (Enderlein); Orian, 1972: 2. [Misspelling.] 

Hostplants. Hibiscus sp., Urena lobata (Malvaceae). 

Recorded distribution. Amboina, Philippines, India, Ceylon, Reunion, Madagascar. 

Mesohomotoma tessmanni (Aulmann) 
(Figs 16, 17, 38, 39) 

Udamostigma tessmanni Aulmann, 1912a: 10; 1912b: 101; 1913: 81. Syntypes d\ 'Spanish Guinea' [not 

examined]. 
Mesohomotoma (Udamostigma) tessmanni (Aulmann) Cotterell, 1927: 109. 
Mesohomotoma tessmanni (Aulmann); Alibert, 1951: 44; Eastop, 1961: 167; Forsyth, 1966: 73; Leston & 

Gibbs, 1968: 73; Roberts, 1969: 76; Leston, 1973: 322. 
Tyora tessmanni (Aulmann) Eastop, 1958: 19; Kaufmann, 1973: 285. 
Tyora (Mesohomotoma) tessmanni (Aulmann); Wood, 1980: 174. 

Hostplants. Cola spp., Theobroma cacao, Sterculia tragacantha (see Forsyth, 1966), Heritiera spp. 
(BMNH) (Sterculiaceae); Desplatzia lutea (BMNH) (Tiliaceae); Chytranthus talbotii (see Forsyth, 1966) 
(Sapindaceae); Cleistopholis sp. (BMNH) (Annonaceae) (possible host misidentification). 

Recorded distribution. Ghana, Nigeria, Equatorial Guinea. 

Material examined 

Numerous adults and larvae from 
Sierra Leone (on Heritiera, Cola, Theobroma), Ivory Coast (on Heritiera), Ghana (on Theobroma, 
Desplatzia), Nigeria (on Cola, Theobroma), San Tome (on Theobroma), Cameroon (on Cola, Theobro- 
ma), Central African Republic (on Cola), Gabon (on Cleistopholis), Uganda (on Cola), Zaire and Angola. 

Comments. Cotterell (1927) records this species as a minor pest of cocoa (Theobroma cacao) in West 
Africa, damaging young shoots and shortening internodes, which causes 'bunching'. Wood (1980) 
summarises that, under conditions of drought and high insolation, heavy infestations cause dessication and 
death of terminal buds. However, Kaufmann (1973) reports that when the insects infest the flower buds 
there is little effect on pod production and the insects may be involved in flower pollination. 

Phylogeny, biogeography and hostplants 

(Summary Fig. 1) 

The Carsidaridae and Homotomidae are considered here to be sister-groups, the pair being 
characterised as follows: antennal sockets enlarged and giving the head a cleft appearance in 
dorsal view; paired metapostnotal epiphyses present; ventral sense organs of hind femora 
situated basally and not arranged linearly. Hostplants of both families are in the Malviflorae 
(sensu Thorne, 1983). Homotomids have a characteristic bipartite male proctiger and many 
have a thickened and densely hirsute antennal flagellum, although the Macrohomotoma-group 
of genera have a simple filiform flagellum. All homotomids develop on species of Ficus 
(Urticales) and are distributed mainly in the Old World tropics, but one small genus occurs in 
Central and northern South America. Two species occur in the Mediterranean Basin on Ficus 
carica, and a third species extends into eastern South Africa on Ficus petersii. Carsidarids are 
diagnosed by the character-suite given on p. 89 and are restricted to hostplants in the Malvales; 
the group is almost entirely pantropical. 

Within the Carsidaridae the Central American genus Epicarsa has a distinctive forewing and 
antennal flagellum; the structure of the male proctiger is unique and the genus is considered to 
be the sister-group of the rest of the family. The single known species is found on Ceiba 
(Bombacaceae). Apart from Epicarsa and the Central and South American genus Paracarsidara 
the rest of the family is distributed in the Old World. Carsidara is quite distinct, having highly 
modified female terminalia and male aedeagal apex; it also retains the primtively broad forewing 
and very short M+Cu stem and has species in tropical Africa, Madagascar and South East Asia 



MALVALES-FEEDING PSYLLIDS 109 

on sterculiaceous hosts. Of the narrow-winged genera Protyora is unique in retaining the 
primitive features of a costal break in the forewing and a simple, flask-shaped male proctiger; it 
is separated from the rest of the group by the derived state of the aedeagal apex. Two species are 
known, one in Sulawesi Utara and another in Australia on a sterculiaceous host. The remaining 
genera are difficult to characterise as a whole; they all possess a male proctiger that is enlarged 
basally and bears small or large lateral lobes but this character is also shared by Carsidara and is 
not likely to be a parallel development in this genus. Within this assemblage Tyora and 
Tenaphalara are grouped together, having an additional rj-rs crossvein and hind tibial spurs 
arranged 1+2+2. Tyora species have retained many primitive features, including the presence 
of radular spinules in cell cuj and the close proximity of the apex of the calval suture to the apex 
of Cu Ib ; they are restricted to sterculiaceous hosts in South East Asia. Tenaphalara has more 
derived features, its species occur in both Africa and the Oriental Region on hostplants in the 
Bombacaceae. Mesohomotoma, with species in the Afrotropical, Malagasy, Oriental, Australa- 
sian and Pacific Regions, has a distinctive bipartite male proctiger that bears a median posterior 
lobe; it also lacks a pterostigma in the forewing but this feature is shared, probably by 
convergence, with some species of Allocarsidara . The form of the endtube of the ductus 
ejaculatorius is similar to that of Paracarsidara and this is considered as a synapomorphy for the 
two genera. Mesohomotoma species usually develop on malvaceous hosts but one African 
species is known to live on various Sterculiaceae. Species of the Central and South American 
genus Paracarsidara develop on both Bombacaceae and Malvaceae. Little is known of the 
hostplants of the South East Asian genus Allocarsidara but three species develop on Durio 
zibethinus (Bombacaceae). The sister-group relationship of Allocarsidara is not resolved. It has 
rounded margins of the vertex, as in Tenaphalara, and some of the species lack a pterostigma, as 
in Mesohomotoma, but both these characters are likely to be parallel developments. 

A list of characters is given below and the matrix, from which the phylogeny (Fig. 1) was 
generated, is given in Table 2. 

Raven & Axelrod (1974), in their review of angiosperm biogeography, state, with reserva- 
tions, that the primary radiation of both the Malvales and the Moraceae occurred in Africa and 
South America in the Upper Cretaceous period. In the Malvales, the Sterculiaceae were the first 
group to reach Eurasia, followed by the Bombacaceae and then the Malvaceae. This is 
reasonably congruent with the hostplant preferences of the carsidarid genera. Those displaying 
more primitive features, e.g. Carsidara, Tyora and Protyora, are on Sterculiaceae, while those 
with more derived features, e.g. Tenaphalara, Allocarsidara, Paracarsidara and Mesohomoto- 
ma, are on Bombacaceae and Malvaceae. Mesohomotoma is anomalous as its species develop 
on both Malvaceae and Sterculiaceae, but the Sterculiaceae-feeding habit is restricted to one 
African Mesohomotoma species and may well be a secondarily derived condition. 

If Fig. 1 is a good reflection of carsidarid phylogeny, the subgroups must have been 
differentiated before the complete separation of Africa from South America (100 m.y.B.P.) as 
elements of the assemblage showing the most derived characters, e.g. Allocarsidara + Paracar- 
sidara + Mesohomotoma, are present today in both continents, and this is unlikely to be through 
subsequent dispersal from West Africa to the Caribbean. However, this is the weakest part of 
the projected phylogeny as these genera are not diagnosed as a group. 

List of characters 

Primitive condition is given in parenthesis 

1. Integument of head and thorax sparsely short-setose or almost glabrous. (Integument of head and 
thorax densely long-setose.) 

2. Flagellomeres thickened. (Flagellomeres narrow, elongate.) 

3. Flagellomeres thickened and densely hirsute. (Falgellomeres narrow, elongate.) 

4. Flagellomere 3 with subapical rhinarium. (Rhinarium absent from flagellomere 3.) 

5. Forewing angular apically. (Forewing rounded apically.) 

6. Forewing narrow. (Forewing broad.) 

7. Costal break absent. (Costal break present.) 

8. Pterostigma absent. (Pterostigma present.) 

9. Rs and M 1+2 anastomosing. (Rs and M 1+] parallel, not in contact.) 



110 D. HOLLIS 



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10. Non-tracheate rs-m crossvein present, (rs-m crossvein absent.) 

11. Non-tracheate rj-rs crossvein present. (r r rs crossvein absent.) 

12. M+Cu stem long. (M+Cu stem short.) 

13. Apical spurs of hind tibia arranged 1+2+2. (Hind tibial spurs arranged l+2(or 3) + l.) 

14. Metabasitarsus with a single apical spur. (Metabasitarsus with two apical spurs.) 

15. Male proctiger bipartite, apical section consisting of a simple anal tube. (Male proctiger unipartite, 
tubular.) 

16. Male proctiger bipartite, apical section having a median posterior lobe in addition to anal tube. (Male 
proctiger unipartite, tubular.) 

17. Male proctiger with enlarged basal portion subdivided posteroapically into small lateral lobes. (Male 
proctiger tubular.) 

18. Male proctiger with well-developed, simple lateral lobes. (Male proctiger tubular.) 

19. Lateral lobes of male proctiger each with a secondary inward-pointing lobe. (Secondary, inward- 
pointing lobes absent.) 

20. Male subgenital plate with supplementary lobes. (Male subgenital plate without supplementary 
lobes.) 

21. Inner lateral surfaces of male subgenital plate with patches of spinules. (These spinules not differenti- 
ated.) 

22. Apex of aedeagus enlarged, endtube of ductus ejaculatorius simple. (Apex of aedeagus narrow, 
endtube of ductus ejaculatorius simple.) 

23. Apex of aedeagus enlarged, complex, endtube of ductus ejaculatorius laterally compressed and 
heavily chitinised. (Apex of aedeagus narrow, endtube of ductus ejaculatorius simple.) 

24. Ductus ejaculatorius with a complex endtube bearing ridges and hooks. (Endtube of ductus ejacula- 
torius without ridges or hooks.) 

25. Female proctiger with a posterodorsal lobe. (Female proctiger without a posterodorsal lobe.) 

26. Lateral margins of female proctiger densely long-setose. (Lateral margins of female proctiger sparsely 
short-setose.) 

27. Lateral palps of ovipositor heavily ridged. (Ovipositor palps not ridged.) 



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MALVALES-FEEDING PSYLLIDS 113 

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1920. The Psyllidae of Borneo. Philippine Journal of Science 17: 353-359. 

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Ferris, G. F. 1928. Observations on the Chermidae (Hemiptera: Homoptera). Part V. Canadian 

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1923. Forest Insects of Australia, viii -I- 171 pp. Sydney. 

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114 D. HOLLIS 

Hollis, D. 1976. Jumping plant lice of the tribe Ciriacremini (Homoptera: Psylloidea) in the Ethiopian 

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MALVALES-FEEDING PSYLLIDS 115 

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Yang, Chung Tu 1984. Psyllidae of Taiwan. Taiwan Museum Special Publication 3: 1-305. 



116 



D. HOLLIS 




Figs 2-17 Carsidaridae, head and antennal structure. 2, Epicarsa corniculata, head and antenna, dorsal 
view. 3,4, Carsidara africana; 3, head, dorsal view; 4, same, lateral view. 5,6, Protyora wilsoni; 5, head, 
dorsal view; 6, same, lateral view. 7,8, Tyora congrua; 7, head, dorsal view; 8, same, lateral view. 9, 10, 
Tenaphalara camerunus; 9, head, dorsal view; 10, same, lateral view. 11-13, Allocarsidara iriana; 11, 
head, dorsal view; 12, same, lateral view; 13, flagellomeres 7 and 8. 14, 15, Paracarsidara gigantea; 14, 
head, dorsal view; 15, same lateral view. 16, 17, Mesohomotoma tessmanni; 16, head, dorsal view; 17, 
same, lateral view. Scale lines: Fig. 13 0-1 mm, remainder 0-5 mm. Setosity not shown. Figs 3-12, 14-17 
drawn from dry-mounted specimens. 



MALVALES-FEEDING PSYLLIDS 



117 







Figs 18-28 Carsidaridae forewings. 18, Epicarsa corniculata. 19, Carsidara africana. 20, 21, Tyora 
congrua; 20, from Sulawesi; 21, from Java. 22, T. buxtoni (Solomon Is). 23, T. ornata. 24, T. striata. 25, 
Tenaphalara acutipennis. 26, T. camerunus. 27, Protyorasterculiae. 28, P. wilsoni. Scale lines, 0-5 mm.' 
Setosity not shown. 



118 



D. HOLLIS 












Figs 29-39 Carsidaridae forewings. 29, Allocarsidara malayensis. 30, A. incognita 31, A. iriana. 32, A. 
Juliana. 33, A. bakeri. 34, A. elongata. 35, Paracarsidara gigantea. 36, 37, Mesohomotoma hibisci, 36 
from Sulawesi; 37, from Seychelles. 38, 39, M. tessmanni; 38, from Angola; 39, from Sierra Leone. Scale 
lines, 0-5 mm. Setosity not shown. 



MALVALES-FEEDING PSYLLIDS 



119 




Figs 40-45 Carsidaridae male genitalia. 40-42, Carsidara africana; 40, proctiger and subgenital plate, 
lateral view; 41 , right paramere, inner lateral view; 42, apical segment of aedeagus, lateral view. 43-45,' 
Protyora wilsoni; 43, proctiger and subgenital plate, lateral view; 44, right paramere, inner lateral view; 
45, apical segment of aedeagus, lateral view. Scale lines, 0-1 mm. 



120 



D. HOLLIS 




Figs 46-56 Carsidaridae male genitalia. 46, 47, Tyora buxtoni (Solomon Is) ; 46, proctiger and subgenital 
plate, lateral view; 47, right paramere, anterolateral view. 48, 49, T. ornata (Fiji); 48, proctiger, lateral 
view; 49, right paramere, anterolateral view. 50, T. congrua (Sulawesi), apical segment of aedeagus, 
lateral view. 51-53, Tenaphalara acutipennis; 51, proctiger and subgenital plate, lateral view; 52, right 
paramere, anterolateral view; 53, apical segment of aedeagus, lateral view. 54-56, T. camerunus; 54, 
proctiger and subgenital plate, lateral view; 55, right paramere, anterolateral view; 56, apical segment of 
aedeagus, lateral view. Scale lines, 0-1 mm. 



MALVALES-FEEDING PSYLLIDS 



121 




Figs 57-65 Carsidaridae male genitalia. 57-59, Allocarsidara malayensis; 57, proctiger and subgenital 
plate, lateral view; 58, right paramere, inner lateral view; 59, apical segment of aedeagus, lateral view. 
60-62, A. elongata; 60, proctiger and subgenital plate, lateral view; 61, right paramere, inner lateral 
view; 62, apical segment of aedeagus, lateral view. 63-65, A. iriana; 63, proctiger and subgenital plate, 
lateral view; 64, right paramere, inner lateral view; 65, apical segment of aedeagus, lateral view. Area of 
differentiated spinules indicated within pecked lines in Figs 57, 60, 63. Scale lines, 0-1 mm. 



122 



D. HOLLIS 





Fies *6-74 Carsidaridae male genitalia. 66-68, Allocarsidara Juliana; 66, proctiger and subgenital I plate, 
fcteral 1 view; 67, right paramere, inner lateral view; 68, apieal segment of aedeagus, lateral view. 69-7 1, 
Abakeri; 69, proctiger and subgenital plate, lateral view; 70 right paramere, inner ateral view 71 
aoical segment of aedeagus, lateral view. 72-74, A. incognita; 72, proctiger and subgenital plate lateral 
view 73 g right paramefe, inner lateral view; 74, apieal segment of aedeagus, later* view. Area of 
differentiated spinules indicated within pecked lines in Figs 66, 69, 72. Scale lines, 01 mm. 



MALVALES-FEEDING PSYLLIDS 



123 






Figs 75-78 Carsidaridae male genitalia. 75, 76, Paracarsidara dugesii; 75, proctiger and subgenital plate, 
lateral view; 76, apical segment of aedeagus, lateral view. 77, 78, Mesohomotoma hibisci (Seychelles); 
77, proctiger and subgenital plate, lateral view; 78, apical segment of aedeagus, lateral view. Scale lines, 
0-1 mm. 



124 



D. HOLLIS 




Figs 79-89 Carsidaridae female terminalia, lateral views. 79, Carsidara africana; 80, Protyora wilsom; 
81 Tyora congrua (Sulawesi); 82, T. striata; 83, Tenaphalara camerunus; 84, Allocarsidara juhana; 85, 
A', incognita; 86, A. malayensis; 87, A. elongata; 88, Paracarsidara dugesii; 89, Mesohomotoma hibisci 
(Tanzania). Scale line, 0-1 mm. 



MALVALES-FEEDING PSYLLIDS 



125 








Figs 90-94 Carsidaridae larval structure, dorsal (d) and ventral (v) views of caudal area showing 
arrangement of pore bands. 90, Protyora sterculiae, also showing chaetotaxy; 91, Tenaphalara camer- 
unus; 92, Allocarsidara incognita; 93, Paracarsidara dugesii; 94, Mesohomotoma hibisci (Tanzania). 
Scale lines, 0-1 mm. 



126 



D. HOLLIS 

Index to hostplants 



Invalid names are in italics. 



Abutilon 106 

indicum 88, 106, 107 
Anacardiaceae 99 
Annonaceae 108 
Aphanamixis grandifolia 99 



Bombacaceae 87, 88, 89, 91, 98, 
99,101,102,104,105,108, 
109, 111 
Bombax 98 

buonopozense 88, 99 

ceiba 88, 98, 99 

cyathophorum 105 

malabaricum 88 

sessile 99 
Boraginaceae 107 
Brachychiton 92, 94 

populneum 88, 95 

rupestris 93 

sp. 95 



Cedrela 90 

Ceiba 91, 98, 104, 108 

burchellii 105 

pentrandra 88, 99 

sp. 92, 105 
Chytranthus talbotii 108 
Cinnamomum camphora 107 
Cleistopholis 108 

sp. 108 
Cola 106, 108 

spp. 108 
Cordia dichotoma 107 



Desplatzia 108 

lutea 108 
Dimocarpus longana 99 
Durio 101 

zibethinus 88, 101,102, 103, 
104, 109 

Ebenaceae 100 

Ficus 90, 108 

carica 108 

petersii 108 

spp. Ill 
Firmiana 92 

simplex 88, 93 

Gossampinus malabarica 99 

Heritiera 96, 106, 108 

littoralis 88, 97 

spp. 108 
Hibiscus 106 

boryanus 107 

esculentus 98 

mutabilis 107 

rosasinensis 107 

sp. 107, 108 

tiliaceus 88, 106, 107 

Lauraceae 107 

Malachra alceifolia 105 
Mallotus japonicus 93 
Malva 104 
sp. 105 



Malvaceae 87, 88, 89, 104, 105, 
106,107,108,109,111 

Malvales 87, 88, 89, 90, 108, 109, 
111 

Malviflorae 87, 108, 111 

Mangifera indica 99 

Meliaceae 90, 99 

Moraceae 109 

Palaquium formosanum 100 
sp. 100 

Sapindaceae 99, 108 
Sapotaceae 100 
Sterculia96, 106 

foetida 98 

lanceolata 97 

sp.97 

tragacantha 108 

ureolata 97 
Sterculiaceae 87, 88, 89, 92, 94, 
96,97,106,107,108,109,111 

Theobroma 106, 108 

cacao 88, 108 
Thespesia 106 

populnea 107 
Tiliaceae 108 

Urena 106 

lobata 106, 107, 108 
Urticales 108, 111 

Wissadula 104 
perplocifolia 105 



Index 

Invalid names are in italics; principal references are in bold. 



Aconopsylla 89 
acutipennis 88, 98-99 
africana, Carsidara 93-94 
africana, Mesohomotoma 106, 107 
Allocarsidara 87, 90, 91 (key), 94, 

96, 100-101, 104, 106, 109, 

110,111 
Aphalaridae 89 
aphanamixis 99 
Apsylla 90 

Baccha 88 

bakeril01,102, 103, 104 
buxtoni 96 

Calophyidae 87, 89, 90, 100 
camerunus, Carsidara 99 



camerunus, Tenaphalara 98, 99 

camphorae 106, 107 

Carsidara 87, 89, 90, 91 (key), 92, 

94,104,106,108,109,110, 

111 
Carsidara sp. 93 
Carsidaridae 87, 88, 89-90, 94, 

100,108,110,111 
Carsidarinae 89, 100 
Carsidaroida 89, 95 
Ciriacreminae 89 
Ciriacremum 90 
concolor, Carsidara 105 
concolor, Paracarsidara 105 
confiuens, Strogylocephala 100 
confluens, Synaphalara 100 
confiuens, Tenaphalara 100 



congrua 95, 96-97 
corniculata, Epicarsa 91, 92 
corniculata, Epiciasa 92 
corniculata, Epicrasa 92 
crigi, Deraeocoris 88 

Deraeocoris 88 
Diceraopsylla 89 
Diclidophlebia 89 
dimocarpi 99 

dugesii, Carsidara 104, 105 
dugesii, Paracarsidara 105 
Dynopsylla 92 

elongata, Allocarsidara 101, 104 
elongata, Ctenophalara 98 
elongata, Tenaphalara 98, 99 



Epicarsa 87, 89, 90 (key), 91-92, 

108, 110, 111 
Epiciasa9\ 

Epicrasa 9 1 
Eustigmia 92 

fascipennis, Strogylocephala 

99-100 
fascipennis, Tenaphalara 98, 99, 

100 

gigantea, Carsidara 105 
gigantea, Paracarsidara 105 
gossampini 99 
guandongana 97 

Haplaphalara 89 

heterocephala, Carsidaroida 95, 96 
heterocephala, Nesiope 96 
hibisci, Mesohomotoma 106, 107 
hibisci, Tyora 106, 107 
hibisci, Udamostigma 107 
Homotomidae 87, 90, 91, 108, 
110, 111 

incognita 101, 102-103 
indica, Tyora 107 
indica, Tyoria 108 
intermedia, Nesiope 96 
irianalOl, 103 

javana, Sphaerophoria 88 
Juliana, Allocarsidara 100, 101, 

102, 103, 104 
Juliana, Tenaphalara 98, 101, 102 

limbata, Carsidara 93 
limbata, Thysanogyna 93 
lineaticollis 106, 107 
lutheri, Mesohomotoma 106, 
107-108 



MALVALES-FEEDING PSYLLIDS 

lutheri, Udamostigma 107 pulchriformis, Baccha 88 



127 



Macrohomotoma 108 
malayensis, Allocarsidara 100, 

101, 102, 103 
malayensis, Tenaphalara 98, 100, 

101, 102 
mangiferae 99 
marginalis92,93,94 
Mastigimas 87, 89, 90 
Mesohomotoma 87, 88, 89, 90 

(key), 92, 94, 96, 101,104, 

106,107,109, 110, 111 
mexicana, Carsidara 105 
mexicana, Paracarsidara 105 
minor, Dynopsylla 92, 93 
minor, Eustigmia 93 
minor, Thysanogyna 93 

Neocarsidara 94, 95 
Nesiope 89,95 

obsoleta-group, Trioza 100 
ornata, Nesiope 95, 97 
ornata, Tyora 96, 97 

Paracarsidara 87, 89, 90, 91 (key), 

92,94,96,101,104,106,108, 

109,110,111 
Phacopteronidae 89, 90 
picta, Baccha 88 
praeusta, Baccha 88 
Prionocnemidinae 89 
Prionocnemina 89 
Prionocneminae 89 
Protyora87,89,90,91(key), 

94-95,109,110,111 
pseudonervosa, Nesiope 99 
pseudonervosa, Tenaphalara 98, 

99 
pseudonervosa, Tyora 99 
Psyllidae 89, 90 



rostrata, Carsidara 105 
rostrata, Paracarsidara 105 

shikokuensis, Carsidara 93 
shikokuensis, Thysanogyna 93 
sinuata, Baccha 88 
Sphaerophoria 88 
sterculiae, Neocarsidara 95 
sterculiae, Protyora 88, 95 
sterculiae, Tyora 94, 95 
striata, Tenaphalara 97, 98 
striata, Tyora 96, 97 
Strogylocephala 87, 89, 100 
sulcata, Tenaphalara 97, 100 
Synaphalara 87, 100 

Tenaphalara 87, 89, 90, 91 (key), 

94,96,97-98,99, 100, 101, 

109,110,111 
Tenaphalarini 89 
tessmanni, Mesohomotoma 88, 

106, 108 
tessmanni, Tyora 108 
tessmanni, Udamostigma 108 
Thysanogyna 92 
Togepsylla 89 
Trioza 100 
Triozidae 88, 89, 90 
triozipennis, Tenaphalara 98, 100 
triozipennis, Trioza 100 
Trisetipsylla 90 
Tyora 87, 89, 90, 91 (key), 94, 

95-96,98,106,109,110,111 
Tyoria 106 

Udamostigma 106 
umalii, Tenaphalara 100 
umalii, Trioza 100 

wilsoni 95 



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The legume-feeding psyllids (Homoptera) of the west Palaearctic Region 

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A review of the Malvales-feeding psy Hid family Carsidaridae (Homoptera) 

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A review of the Rhadalinae (= Aplocneminae) (Coleoptera: Melyridae) 

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A review of the Rhadalinae 
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Enid R. Peacock 



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17 December 1987 



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Entomology series 
Vol 56 No 3 pp 129-170 



Issued 17 December 1987 



A review of the Rhadalinae (= Aplocneminae) 
(Coleoptera: Melyridae) 

Enid R. Peacock 

Department of Entomology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Contents 

Synopsis 129 

Introduction 129 

Melyridae Leach, 1815 130 

Distinguishing features 130 

Rhadalinae LeConte 131 

Taxonomy and distinguishing features 131 

Adults 131 

Larvae 131 

Biology 131 

Checklist of the genera of the Rhadalinae 132 

Notes on the key and checklists of species 133 

Abbreviations 133 

Key to adults of the world genera of the Rhadalinae 133 

Generic diagnoses 144 

Acknowledgements 162 

References 162 

Index 167 

Synopsis 

Thirteen world genera of Rhadalinae (Melyridae) are recognized here, of which three are included in the 
subfamily for the first time: Antinea Peyerimhoff, Indiodasytes Pic and Hemipleurus gen. n. , erected for H. 
floriger sp. n. from Borneo. A generic key is given together with a diagnosis and type species citation for 
each genus. Eurelymis Casey (NW. America), from the Melyrinae, and Celsus Lewis (Japan), from the 
Dasytinae, are regarded as junior synonyms of Semijulistus Schilsky (Holarctic Region), Cymbolus 
Gorham is synonymized with Rhadalus LeConte and Xamerpus Fairmaire with Malthacodes Waterhouse. 
The two Indian species of Aplocnemus Stephens, A. indicus Champion and A. moestus Gorham, are 
transferred to Malthacodes Waterhouse. Malthacodes parvus nom. n. is proposed as a replacement name 
for M. minutus (Pic, 1931) [originally in Xamerpus] (nee Pic, 1906). Checklists of genera and species are 
provided. Distinguishing features are given for adult and larval Melyridae and Rhadalinae and comments 
are made on the biology of the latter. 

Introduction 

The Melyridae (sensu Crowson, 1964 and Lawrence, 1982) is the largest family in the 
superfamily Cleroidea, with at least 200 genera. Crowson (1964) considered it to be divisible into 
five subfamilies: the Rhadalinae (Aplocneminae), the Melyrinae, the Malachiinae, the 
Prionocerinae and the Dasytinae. These subfamilies have all, at one time or another, been given 
family rank and some are currently considered as such by recent authors (e.g. Constantin, 1965; 
Liberti, 1984; Wittmer, 1984 etc.). In spite of its size the family has been poorly studied in recent 
years, except for the subfamily Malachiinae which has been the subject of extensive studies by 
Wittmer (1930 et seq.). At the beginning of the century and before, there were several prolific 
writers whose unco-ordinated descriptive efforts, mainly at specific level, caused a certain 
amount of taxonomic confusion in the family. 



Bull. Br. Mus. nat. Hist. (Ent.) 56 (3): 129-170 Issued 17 December 1987 



130 E. R. PEACOCK 

The first author to create some order out of this confusion was Crowson who, in his major 
work on the higher classification of Coleoptera (1955) and in subsequent works (1964, 1970), 
keyed out and defined the superfamilies, families and subfamilies. Since then, Majer (1987) has 
produced a useful work completely revising the phylogeny and taxonomy of the family. He 
introduced new taxa, divided the family into subfamilies, supertribes and tribes and discussed in 
detail the supposed phylogeny of the group. 

In the course of his study of the Melyridae, Crowson (1964) discovered a distinctive group of 
genera whose close relationship had been hitherto overlooked and for which he erected a new 
subfamily, the Aplocneminae. This included genera from the melyrid subfamilies Melyrinae and 
Dasytinae, the Cleridae and the Rhadalidae (Coleopterorum Catalogus, Pic, 1929, 1937; 
Corporaal, 1950; Pic, 1926). However, Crowson overlooked the fact that a family group name 
(Rhadalidae) was already in use for one of his included genera: Rhadalus LeConte. The name 
Rhadalinae therefore has priority (Article 23(a), International Code of Zoological Nomencla- 
ture, 1985) and is employed here. 

These discoveries, along with some additional genera, nomenclatural changes, new generic 
synonymies and the erection of a new genus, indicated the need for a review of the entire 
subfamily and hence this study. 

In the present paper Crowson's (1964) treatment of the subfamily is reappraised, decisions 
concerning the status of the included genera being based on the study of their type species 
(something that Crowson failed to do). As a result 12 of the 25 nominal genera are regarded as 
valid and one additional genus is described as new. 

Although the Rhadalinae comprises only 13 (out of 25 nominal) genera, its distribution is 
almost world-wide. The group seems to be restricted mainly to semi-dry forested, mountainous 
or hilly areas, although some species have been collected on sea beaches. The adult beetles are 
found on flowers, shrubs or on the leaves or bark of coniferous or deciduous trees and although 
most are carnivorous to a certain extent, some are pollinivorous or both carnivorous and 
pollinivorous. The subfamily is well defined, all of its species exhibiting the unusual feature of 
the connation of the first two visible abdominal sternites. This feature is found in no other group 
of Melyridae nor in other cleroid families. Its other unusual and notable feature is the sclerite 
articulated to the (morphologically) dorsal side of the median lobe of the aedeagus. This sclerite 
articulates at its basal end with the median lobe in a hinge-like joint, and is freely moveable in all 
the genera examined except Microjulistus , in which it appears to be fused at the base to the 
median lobe. The only genus in the group lacking this sclerite (or appendage) is the Indian 
Indiodasytes . 

MELYRIDAE Leach, 1815 1 

Distinguishing features 

Adults. i\dult Melyridae are generally distinguished from related families by the following characters: 
antennae usually 11-segmented and serrate or filiform, but sometimes pectinate or with some basal 
segments modified, apical segments not differentiated to form a club; claws either simple, split, toothed or 
with membranous appendages; front coxal cavities open behind, coxae large and projecting, with exposed 
trochantins; tarsi 5-segmented, sometimes 4,5,5 in male [exception Anthriboclerus , 4,4,4] and without 
strongly lobed segments; prothorax usually with distinct side margins; elytra without striae (except in some 
Melyris spp. which also have costae); sometimes with exsertile vesicles at anterior angles of prothorax and 
abdominal pleura; abdomen with 6 visible sternites which are usually free but occasionally with the first two 
connate; aedeagus with undivided tegmen. 

Larvae. Melyrid larvae can be distinguished from those of all other cleroid families [except Phycosecidae] 
by the following characters: head with a well-marked median epicranial suture and no endocarina; 
mouthparts retracted, stipes much longer than cardo; spiracles annuliform (Crowson, 1970); mandible 
with a long, stiff prosthecal process near the middle or at the base of inner margin (Boving & Craighead, 
1930) . They differ from Phycosecidae in possessing 1-5 ocelli on each side instead of 6. [The larvae of other 
cleroid families have the following characters: head rarely with a distinct median cranial suture, but if so, 

1 Fide Watt (1975: 33). Pic (1937: 3) and Majer (1987: 784) give Olivier, 1790 as author but there is no family group 
name in that work. 



REVIEW OF THE RHADALINAE 131 

then with mouthparts strongly protracted and stipes not longer than cardo; endocarina usually distinct; 
spiracles usually bicameral; mandible either without a prosthecal process or with a short one.] 

RHADALINAE LeConte 

Rhadalini LeConte, 1861: 191, 194; LeConte & Horn, 1883: 213, 216. 

'Haplocnemates' Mulsant & Rey, 1868: 181. 

Rhadalinae LeConte; Casey, 1895: 457; Blaisdell, 1938: 3; Hatch, 1962: 86; Arnett, 1968: 609, 611, 613; 

Blackwelder, 1975: R67.10. 
Rhadalidae LeConte; Pic, 1926a: 3; Crowson, 1964: 315, 320. 
Aplocneminae {[erratim Haplocneminae] Crowson, 1964: 316, 317, 318, 319, 320; Constantin, 1965: 92; 

Vinson, 1967: 330); Majer, 1983: 387. Syn. n. 
Aplocnemina; Majer, 1987: 800. 

Taxonomy and distinguishing features 

Adults Adult Rhadalinae are distinguished from the other subfamilies of Melyridae by the following 
characters in combination:- first two visible abdominal sternites connate; apical segment of maxillary palps 
broadened, securiform or triangular except in some species of Aplocnemus Stephens); median lobe of 
aedeagus with dorsal appendage (Fig. 7) or 'lever' (Majer, 1982). 

Members of the Rhadalinae are diverse in appearance but all have the three characters mentioned 
above, except for Aplocnemus (Ischnopalpus) which has a spindle-shaped apical segment to the maxillary 
palps, and Indiodasytes which lacks the dorsal appendage to the median lobe of the aedeagus. They are also 
characterized by the following features:- antennae usually serrate, sometimes pectinate; eyes entire or 
weakly emarginate; head generally broader than long, often retractable under pronotum; body usually 
densely hairy and strongly convex; tarsi varying from finely elongate and simple to short, broad and weakly 
lobed, segment 1 normally at least as long as 2; tarsal claws toothed or with free membranous appendages; 
dorsum either unicolorous or patterned; puncturation on head simple, rimmed, or confused; pronotal 
punctures entirely simple, or simple on disc but rimmed or tuberculate at sides, or all rimmed or all 
tuberculate, or confused so that individual punctures are not distinguishable; elytral punctures usually 
simple, sometimes with a seta emerging from the puncture, but more usually from beside it; epipleura well 
developed at base; wing venation variable as shown in Figs 48-59. 

Crowson (1964) erected the subfamily Aplocneminae (as Haplocneminae) [= Rhadalinae] for the 
following genera: Anthriboclerus Schenkling, [H] aplocnemus Stephens, Cymbolus Gorham, Diplambe 
Schilsky, Donaldia Alluaud, Eucymbolus Champion, Ischnopalpus Schilsky, Julistus Kiesenwetter, 
Malthacodes Waterhouse, Pelecophora Dejean, Rhadalus LeConte, Trichoceble Thomson and Xamerpus 
Fairmaire. Majer (1983), who regards the group as of lower taxonomic rank than subfamily, added 
Kubanius Majer, Semijulistus Schilsky and Microjulistus Reitter. In this paper I add: Antinea Peyerimhoff , 
Celsus Lewis, Eurelymis Casey, Indiodasytes Pic and Hemipleurus gen. n. A complete checklist of included 
genera is provided below. 

Larvae. The known rhadaline larvae {Trichoceble, Aplocnemus, and Pelecophora) possess the following 
characters (Crowson, 1964): head with 2 ocelli on each side, the larger one being anterior to the smaller, so 
that a line through the 2 would pass through the antennal foramen; urogomphi, on abdominal tergite 9, 
varying from minute to very long; abdominal tergite 9 either with a membranous appendage in the middle 
of each side {Pelecophora and Aplocnemus), or with a group of setiferous tubercles (unnamed larva from 
Chile) or forming a large flat plate with minute, widely separated urogomphi {Trichoceble). 

Rhadaline and melyrine larvae have glandular openings on the 9th abdominal tergite only (Crowson, 
1981) and some Aplocnemus larvae have dark subcuticular patches on abdominal segments 1-8, usually 2 
pairs on each segment. Vinson gives a detailed description of a larva assumed to be Pelecophora pikei 
Vinson (1957). 

Larvae of the other melyrid subfamilies usually have more ocelli, but if there are only two on each side, 
then they are one above the other so that a line through them would not pass through the antennal foramen. 
Tergite 9 is without the above mentioned structures. 

Biology 

With the exception of Australasia, species of Rhadalinae are known to occur in all major biogeographic 
regions, but little is known about their biology. 

Most adult rhadalines are probably carnivorous to a certain extent. Examination of the gut contents of 
adult species of Aplocnemus , Trichoceble, Pelecophora and Donaldia [= Malthacodes] has revealed insect 



132 E. R. PEACOCK 

fragments (Vinson, 1946; Crowson, 1964), but in species of Rhadalus and Indiodasytes only pollen grains 
were found (Crowson, 1964; Peacock, pers. obs.). Denticulation of the cutting edge of the mandible, a 
character indicating pollinivorous habits, is absent in most Rhadalinae, although it is shown in a figure of a 
mandible of Pelecophora (Vinson, 1946), indicating perhaps that this genus is both carnivorous and 
pollinivorous. 

It is likely that the larvae of this group are carnivorous as are most cleroid larvae, even though the 
examination of some larval gut contents by Crowson (1964) revealed no identifiable remains. This view is 
supported by the presence of a pedunculate seta (Vinson called it a 'lacinia') on the maxillary mala of a 
larva of Pelecophora (Vinson, 1957), a character peculiar to all known carnivorous cleroid larvae 
(Crowson, 1964). 

There appear to be scant records of habitat preference and the few larvae that have been collected seem 
to live concealed either on or in the soil or in decaying wood. The following paragraphs give an indication of 
the habitat preferred by some adults. 

In Mauritius, Pelecophora and Donaldia [= Malthacodes] have been found during the day on the leaves 
and branches of shrubs and trees, where they were probably feeding on small insects (Vinson, 1946). 
Vinson noticed that these seemingly carnivorous beetles are constantly associated with the indigenous 
vegetation, occurring in their greatest numbers during the hot season from November to March. He also 
found a Pelecophora larva under a stone in association with an adult crawling out of a pupal cell (1957). 

Adults of the North American Eurelymis [= Semijulistus] are often found at high altitudes. They are 
attracted to wood smoke (Leech, 1931) and have been found on Achillea (Hatch, 1961). E. [= S.] atra 
LeConte has been reared from pupae which were found in bee burrows, in sand, in Alberta, Canada, and 
E. [= S.]flavipes LeConte was found on Pyrocantha in California [specimens loaned to author]. Cymbolus 
[= Rhadalus] elongatus Champion was collected in Temascaltepec, Mexico, in large numbers on Mimosa 
and Spondias spp. and several individuals of C. [= R.] wolcotti Hinton were collected on Pinus 
pseudostrobus Lindley (Hinton, 1934). 

In Japan, Celsus [= Semijulistus] is not common and has only been seen in recent years (Nakane, pers. 
comm.). In Kogoshima, Nakane found adults in abundance on flowers of Viburnum, Rosa?, Prunusl etc. 
in April and May, but collected only one in July. They were usually captured singly or in pairs except in 
mid- April when as many as 45, of both sexes, were seen at a time. No larvae were found. 

In Central Europe Aplocnemus is found on the flowers of pine trees, but has also been collected on 
deciduous trees, and Trichoceble (widespread but not common) has been found on flowering shrubs and in 
dry wood under trees (Lohse, 1979). Aplocnemus has been recorded on oak in Spain (Constantin, 1965) 
and Kubanius elegans Majer (U.S.S.R.) was 'reared in Picea schrenckiana' (Majer, 1983). 

Checklist of the genera of the Rhadalinae 

ANTHRIBOCLER l/SSchenkling, 1922: 328. 
ANTINEA Peyerimhoff , 1929: 191. 
APLOCNEMUS Stephens, 1830: 316. 
Elicopis Stephens, 1829: 136. 
Haplocnemus Stephens; Agassiz, 1846 [7]: 29. 
Helicopis Stephens; Agassiz, 1846[7]: 29. 
Subgenus APLOCNEMUS Stephens, 1830: 316. 
Subgenus DIPLAMBE Schilsky, 1894: 234. 
Subgenus HOLCOPLEURA Schilsky, 1894: 234. 
Subgenus ISCHNOPALPUS Schilsky, 1894: 235. 
Subgenus PSEUDAPHYCTUS Pic, 1896: 47. 
EUCYMBOLUS Champion, 1913: 129. 
HEMIPLEURUS gen. n. 
INDIODASYTESPic, 1916: 14. 
KUBANIUS Majer, 1983: 385. 
MALTHACODES Waterhouse, 1876: 116, 
Xamerpus Fairmaire, 1886: 41. Syn. n. 
Donaldia Alluaud, 1898: 102. 
MICRO JULISTUS Reitter, 1889a: 111. 

Ceralliscus Bourgeois, 1894: 121. 
PELECOPHORA Dejean, 1821: 115. 
Diglobicerus Latreille, 1829: 475. 
RHADALUSLeConte, 1852: 212. 

Cymbolus Gorham, 1886: 324. Syn. n. 



REVIEW OF THE RHADALINAE 133 

SEMIJULISTUS Schilsky, 1894: 227. 

Celsus Lewis, 1895: 118. Syn. n. 

Eurelymis Casey, 1895: 600. Syn. n. 
TRICHOCEBLE Thomson, 1859: 109. 

Julistus Kiesenwetter, 1859: 175. 

Notes on the key and checklists of species 

The measurements of approximate body length in the descriptions are taken from the front of the 
pronotum to the apex of the elytra, since in many genera the head is retractable under the pronotum and 
hence only partially visible from above. 

The term 'free' when describing the membranous appendages on the claws means that they are attached 
to the claws at the base only (Fig. 15). In most dasytines possessing these structures, the appendage is 
attached to the claw along its length almost to the apex, or for at least half its length (Blaisdell, 1938: pi. 2, 
figs 1-20). 

A 'rimmed' puncture means that there is a circle around the puncture but the area within this is not raised 
above the surrounding surface (Fig. 1). A 'tuberculate' puncture is within a circular area which is raised 
above the surrounding surface (Figs 21, 22, 24). 

In the species checklists it should be noted that the species of earlier authors are taken from the 
Coleopterorum Catalogus (Pic, 1926, 1929, 1937) in order to complete the lists and have not in all cases 
been examined. However, the references have been checked and many errors have been rectified. The 
varieties cited in Pic's catalogue have been listed here in synonymy. The generic names in parentheses 
denote the genus in which the species was described, if not the current genus. 

Abbreviations 

BMNH British Museum (Natural History), London. 

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. 

MNHN Museum national d'Histoire naturelle, Paris. 

USNM National Museum of Natural History [U.S. National Museum], Smithsonian Institution, 

Washington, D.C. 
[Note. See Acknowledgements for full list of institutions from which material was borrowed.] 

Key to adults of the world genera of the Rhadalinae 

1 Epipleura very narrow, usually mat or granular, and deflecting laterad at, or before, level of 

hind coxae (Figs 3,4), becoming subvertical. Pubescence of dorsum uniformly short and 
recumbent, sometimes scale-like (Figs 1,2). Tarsi elongate and slender (Figs 3, 5), claws 
with small teeth, without membranous appendages. Antennae fairly short, sometimes with 
segments 7-10 enlarged, compactly serrate (Fig. 1). Length not more than 5 mm 2 

- Epipleura broader, usually shiny, deflecting dorsad either at about level of 2nd or 3rd visible 

abdominal sternite (Figs 8, 17, 35) or at or before base of 1st visible abdominal sternite (Fig. 
9). Pubescence of dorsum not as above, nearly always with some erect or semi-erect setae. 
Tarsi variable. Antennae elongate, reaching base of pronotum or beyond and usually loosely 
serrate or pectinate from segments 4-10 (rarely 3-10) (Figs 40-44) . Length 2-7mm 3 

2 Epipleura evanescent anterior to 1st abdominal sternite (Fig. 4). Lateral pronotal margin 

smooth, or with a few small teeth near base. Pronotum mat, puncturation indistinct, with 
strong microsculpture, forming an irregular reticulum under pubescence (Fig. 2). Disc of 
metasternum flat, with distinct median longitudinal groove (Fig. 4). Females without 
unpunctured patch on apical third of elytron. Length less than 3 mm. (C. Europe & Asia, 
Africa, Arabia, Mediterranean) MICRO JULISTUS (p. 161) 

- Epipleura narrow and becoming subvertical from level of 1st abdominal sternite , visible in side 

view from just behind shoulder. Lateral pronotal margin strongly serrate. Pronotum with 
shallow, rimmed punctures (Fig. 1), sometimes obscured by pubescence. Disc of metaster- 
num convex, without a distinct median longitudinal groove (Fig. 3). Females usually with 
shiny unpunctured patch on apical third of each elytron (Fig. 6). Length 3-5 mm. (C. & E. 
Europe, Asia, Japan, western U.S.A., Canada) SEMIJULISTUS (p. 159) 

3 Tarsi elongate and claws appendiculate, membranous appendages either absent or minute. 

(Figs 10-13) 4 



134 



E. R. PEACOCK 





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Figs 1-7 1, 6, 7, Semijulistus ater (Canada): (1) pronotum; (6) elytral apices of female; (7) aedeagus of 
male. 2, 4, Micro julistus fulvus (U.S.S.R., Kazakhstan): (2) pronotum; (4) metasternum. 3, 5, 
Semijulistus spectabilis (Japan): (3) metasternum; (5) hind tarsus. (Scale line = 0-3 mm.) da = dorsal 
appendage; ml = median lobe.) 



REVIEW OF THE RHADALINAE 



135 





Figs 8-15 8, Malthacodes pictus (Rodriguez I.): epipleuron. 9, Hemipleurus floriger (Sarawak [para- 
type]): epipleuron. 10, 11, Kubanius elegans (U.S.S.R., Kazakhstan [paratype]): (10) hind tarsus; (11) 
hind claw. 12, 13, Trichoceble floralis (France): (12) hind tarsus; (13) claw. 14, Aplocnemus impressus 
(England): hind claw. 15, Pelecophora illigeri, (Mauritius): hind claw. (Scale line = 0-1 mm.) ma = 
membranous appendage; ac = appendiculate claw. 



136 E. R. PEACOCK 

- Tarsi stout or delicate, claws not appendiculate, but with large free membranous appendages 

(Figs 14, 15, 45-47) 6 

4 Eyes small, oval, not projecting beyond contour of head. Head weakly convex between eyes. 

Temples long, subparallel. Claws with minute teeth. Apterous, without humeral swellings. 
(N.Africa: Sahara.) ANTINEA{p. 159) 

- Eyes large, reniform, prominent, projecting beyond contour of head when seen from above. 

Head sometimes with a short, longitudinal groove or depression at inner side of antenna and 
eye. Temples constricted behind eyes. Claws with strong teeth. Winged, elytra with distinct 
humeral swellings 5 

5 Dorsum without metallic sheen. Hard-bodied. Pronotum usually elliptical, broader than head 

(including eyes). Elytra and pronotum convex and densely punctured; epipleura long, 
extending to about 2nd visible abdominal sternite. Hind tarsus not quite as long as tibia. 
Claws with either pointed (Figs 12, 13) or blunt teeth. (C. , S. & E. Europe, Asia, China.) 

TRICHOCEBLE(p. 158) 

- Dorsum with slight metallic sheen. Soft-bodied, malacodermiform. Pronotum subquadrate or 

elliptical, sometimes flattened on disc, not always broader than head. Elytra flattened on 
disc , apices abruptly declivous , rather sparsely punctured ; epipleura short , not reaching hind 
coxae. Hind tarsus about as long as tibia (Fig. 10). Claws with large, blunt teeth, sometimes 
also with small, obscure membranous appendages (Fig. 11). (U.S.S.R. (Kazakhstan), N. 
India.) KUB ANIUS (p .157) 

6 Elytra with sublateral carina in basal half, forming a double elytral edge (Fig. 16) 7 

- Elytra without sublateral carina in basal half (Fig. 17) 9 

7 Pronotal punctures all simple. Apical segment of maxillary palps only weakly securiform (Fig. 

31). Form elongate, elytra more than twice as long as wide. Dorsum usually black, 
sometimes with a metallic sheen. (Greece.) APLOCNEMUS(DIPLAMBE)(p. 151) 

- Pronotal punctures tuberculate, at least near sides. Apical segment of maxillary palps 

triangular. Form elongate and dorsum usually unicolorous brown, or form very broad 
(length less than twice width) , head brown , elytra and pronotum black 8 

8 Form elongate. Elytra and pronotum brown. Scutellum mat, densely and finely punctate and 

densely setose (Fig. 18). Sublateral elytral carina close and subparallel to costal margin, 
forming an abrupt, narrow, horizontal shelf, with a row of very large punctures along inner 
side (Fig. 16). Disc of pronotum with strongly rimmed or tuberculate punctures, sometimes 
with raised, shiny puncture-free patches (Figs 21, 22). (Southern U.S.A., C. & S. America.) 

RHADALUS(p. 144) 

- Form very short and broad (see couplet 7) . Elytra and pronotum bluish-black . Scutellum shiny , 

very sparsely punctate and setose. Sublateral carina oblique to costal margin and less 
prominent, so scarcely forming a shelf, without such large punctures along inner side. 
Pronotum with small simple punctures on disc and tuberculate punctures near sides. (C. 
America.) EUCYMBOL US(p. 145) 

9 Humeral swellings very prominent and elytra with a pronounced transverse depression across 

basal third (Figs 29, 30) 10 

- Humeral swellings less prominent and elytra without a transverse depression across basal third . 

11 

10 Pronotum very large and quadrate, about half as long as elytra; shiny, with sparse setae, 

puncturation inconspicuous (Fig. 27). Head large, shiny and convex with faint diagonal 
ridges from base of antenna to eye (Fig. 25). Elytra with semi-erect shorter setae between the 
sparse, very long, erect setae; tufts of dense setae present on weak tubercles, just posterior to 
transverse elytral depression, and also on larger tubercles at inner side of humeral swellings 
(Fig . 29) ; epipleura extending to base of 2nd visible abdominal sternite . (Seychelles . ) 

ANTHRIBOCLERUS(p. 145) 

- Pronotum of moderate size, transverse, much less than half as long as elytra; with distinct 

puncturation (Fig. 28). Head mat, without diagonal ridges (Fig. 26). Elytra with recumbent 
setae between sparse, long, erect setae (Fig. 30); epipleura not extending further than base 
of 1st visible abdominal sternite (Fig. 9). (Borneo.) HEMIPLEURUS(p. 147) 

11 Pronotal punctures simple (Fig. 19). Maxillary palps moderately securiform (Fig. 31) or 

spindle-shaped (Fig. 32). Dorsum usually brown or black, often with a metallic bluish or 
greenish tinge (some species from the Canary Is are light brown or yellow). Setae on elytra 
usually of fairly uniform length and colour. (Palaearctic, N. & W. Africa (Angola? & Ivory 
Coast), Asia Minor, Philippines.) APLOCNEMUS (p. 151) 



REVIEW OF THE RHADALINAE 



137 




Figs 16-18 16, Rhadalus rufopiceus (Guatemala [syntype]): lateral view of elytron. 17, Indiodasytes 
madurensis (India [holotype]): lateral view of body. 18, Rhadalus rufopiceus: scutellum. (Scale line = 
0-4 mm.) 



138 



E. R. PEACOCK 




Figs 19-24 Pronotum of: (19) Aplocnemus impressus (England); (20) Pelecophora illigeri (Mauritius); 
(21) Rhadalus rufopiceus (Guatemala [syntype]); (22) Rhadalus testaceus (U.S.A.); (23) Trichoceble 
floralis (France); (24) Indiodasytes madurensis (India [holotype]). (Scale line = 0-5 mm.) 



REVIEW OF THE RHADALINAE 



139 




Figs 25-30 25, 27, 29, Anthriboclerus scotti (Seychelles [paratype]); (25) part of head; (27) pronotum; 
(29) basal half of elytra. 26, 28, 30, ditto, Hemipleurus floriger (Sarawak [paratype]). (Scale line = 0-1 
mm in Firs 25, 26; = 0-3 mm in Figs 27-30.) dl = diagonal line. 



140 



E. R. PEACOCK 




Figs 31-35 31-33, apical segment of maxillary palp of: (31) Aplocnemus nigricornis (England); (32) 
Aplocnemus (Ischnopalpus) sanctus (Palestine); (33) Pelecophora illigeri (Mauritius). 34, 35, ventral 
view of prosternum of: (34) Pelecophora illigeri', (35) Malthacodes vageguttatus (Madagascar). (Scale 
line = 0-1 mm in Figs 31-33; = 0-5 mm in Figs 34-35.) 



- Pronotal punctures tuberculate or rimmed at sides. Maxillary palps broadly triangular (Fig. 

33). Dorsum brown or black or bicolorous and patterned. Elytral setae not usually uniform, 
sometimes both erect and semi-erect , or with patches of recumbent setae , often bicolorous . 12 

12 Head very large, not retractable under pronotum. Prosternum long in front of coxae; 
prosternal process well developed (Fig. 34). Pronotal punctures usually simple on disc and 
tuberculate at sides, sometimes becoming weakly tuberculate on disc also, which then 
appears rugose (Fig. 20). Elytra usually with short and long, erect or suberect setae, often 
forming patches. Cuticle or setae bicolorous. (Mauritius, Reunion, E. Africa.) 

PELECOPHORA(p. 146) 

- Head smaller, somewhat vertical, partly retractable under pronotum. Prosternum shorter in 

front of coxae; prosternal process small and pointed (Fig. 35). Pronotal punctures rimmed or 
tuberculate on disc, or not clearly defined, but never simple. Cuticle either bi- or unicolor- 
ous; setae variable in length, type and colour 13 



REVIEW OF THE RHADALINAE 



141 




• 



• 





'::';,- 


e I ', 


■ I^H 


"1 


III 


[38J 


w/M 








Figs 36-39 Elytral puncturation of: (36) Rhadalus rufopiceus (Guatemala); (37) Hemipleurus floriger 
(Sarawak); (38) Semijulistus ater (Canada); (39) Trichoceble floralis (France). (Scale line = 0-1 mm.) 



13 Tarsi stout, hind ones longer than half a tibia, dark brown, without very long ventral setae, 

segments 1-4 subequal, penultimate segment apically truncate dorsally (Fig. 45). Form 
elongate. Dark brown and/or black 14 

- Tarsi delicate , hind ones usually about half tibial length or less , usually pale coloured , segments 

1-3 subequal sometimes with very long ventral setae, segment 4 smaller, dorsally apically 
emarginate (Fig. 46). Form either broad and short or elongate. Unicolorous or bicolorous, 
often patterned. (Madagascar, Seychelles, Aldabra, Mauritius, Reunion, Rodriguez I., 
Africa, India, Sri Lanka.) MALTHACODES [in part] (p. 149) 

14 Head and pronotum black, elytra dark brown. Pronotum very strongly punctured and with a 

distinct longitudinal unpunctured area (sometimes raised) in mid line near base. Dorsal 
pubescence recumbent or semi-erect, usually with a few erect setae laterally (Fig. 24). Wings 
with strong venation, anal cell either present or obsolete. Aedeagus without dorsal appen- 
dage. (India, Sri Lanka.) INDIODASYTES(p. 151) 

- Dorsum entirely dark brown or black. Pronotum less strongly punctured and without longitu- 

dinal unpunctured area near base. Dorsal pubescence not recumbent. Wings with weak 
venation, anal cell lacking. Aedeagus with dorsal appendage. (India.) 

MALTHACODES [in part] (p. 149) 



142 



E. R. PEACOCK 




Figs 40-47 40-44, antenna of: (40) Kubanius elegans (U.S.S.R.: Kazakhstan [paratype]); (41) Malth- 
acodes pictus (Rodriguez I.); (42) Malthacodes vageguttatus (Madagascar); (43) Pelecophora illigeri 
(Mauritius); (44) Hemipleurus floriger (Sarawak [paratype]). 45-47, tarsus of: (45) Indiodasytes 
madurensis (India [holotype]) (hind); (46) Malthacodes vageguttatus (hind); (47) Hemipleurus floriger 
(front). (Scale line = 0-5 mm in Figs 40-44; = 0-1 mm in Figs 45-47.) 



REVIEW OF THE RHADALINAE 



143 




Figs 48-59 Hind wing of: (48) Rhadalus elongatus (Mexico); (49) Anthriboclerm scotti (Seychelles 
[paratype]); (50) Pelecophora illigeri (Mauritius); (51) Hemipleurus floriger (Sarawak [paratype]); (52) 
Malthacodes pictus (Rodriguez I.); (53) Malthacodes vageguttatus (Madagascar); (54) Malthacodes 
elegans (Mauritius); (55) Indiodasytes sp. (undescribed sp. from Sri Lanka); (56) Aplocnemus nigricor- 
nis (France); (57) Trichoceble floralis (France); (58) Semijulistus flavipes (U.S.A., California); (59) 
Microjulistus subconvexus (South Africa). (Scale line = 1-0 mm.) ac = anal cell. 



144 E. R. PEACOCK 

Generic diagnoses 
RHADAL US LeConte 

(Figs 16, 18,21,22,36,48) 

Rhadalus LeConte, 1852: 212. Type species: Rhadalus testaceus LeConte, 1852: 212, by monotypy. 
Cymbolus Gorham, 1886: 324. Type species: Cymbolus rufopiceus Gorham, 1886: 324, by subsequent 
designation (Blaisdell, 1938: 9). Syn.n. 

Elongate, convex, light or dark brown, occasionally darker ventrally or legs, palps and antennae lighter. 
Dorsum and legs covered with long erect, semi-erect or strongly curved golden setae. 

Head subvertical , usually with a longitudinal groove or ridge at each side of f rons , near inner side of eye ; 
puncturation sparse, simple, rimmed or weakly tuberculate, surface shiny or microreticulate between 
punctures. Eyes large or small, varying in shape from entire to strongly emarginate, with short setae; facets 
very large and convex. Antennae long, extending well beyond base of pronotum, length from less than 
two-fifths of body length to more than half; serrate from 4th segment. Apical segment of maxillary palps 
triangular. Pronotum weakly to strongly transverse (Figs 21, 22); lateral margins explanate, curved and 
weakly crenulate; base strongly sinuate, bordered; all angles more or less rounded; either evenly elliptical 
or with anterior angles weakly produced anteriad; disc sometimes with raised shiny patches between 
punctures; punctures usually rimmed or tuberculate on disc, rarely simple, sometimes tuberculate near 
sides; anterior margin of pronotum strongly anterior to that of prosternum, allowing head to fit in 
vertically. Scutellum broadly triangular, with rounded apex, mat (owing to dense puncturation), densely 
setose, cuticle obscured (Fig. 18). Elytral sides parallel or weakly diverging posteriorly, crenulate, with a 
carina running subparallel to lateral edge, becoming obsolete well before apex; above carina a narrow 
horizontal shelf with row of very large punctures in a concavity (Fig. 16); elsewhere with smaller, dense 
punctures, from which elytral setae emerge (Fig. 36); epipleura quite broad, transversely concave, with 
well-defined smooth inner edge , deflecting dorsad and becoming obsolete at about level of 3rd to 5th visible 
abdominal sternite. Hind wing as in Fig. 48. Tarsi moderately elongate with segments 1-4 weakly lobed 
and densely ventrally setose, segment 4 marginally smaller than 3; apical segment almost as long as the 
others together and much broader at apex. Claws with large, free, membranous appendages. Anterior 
tibiae with a few short, rather obscure external spines among long erect and semi-erect setae. Prosternal 
process on higher plane than front coxae, very narrow, just reaching their hind margin. Length 3-8-7-0 mm 
(width 1-7-3-0 mm). 

Distribution. Guatemala, Mexico, Brazil, southern U.S.A. (Arizona). 

Material examined 

8 species, including syntypes of type species of Cymbolus (BMNH) and holotype of type species of 

Rhadalus (MCZ). 

Remarks This genus is recognized by its colour, sublateral elytral carina, strongly setose scutellum, the 
form of the tarsi, the rimmed or tuberculate pronotal punctures and the raised puncture-free pronotal 
patches which are present on the disc of most species. It is also recognized by the fact that the elytral setae 
emerge from the large punctures (Fig. 36) instead of from a smaller adjacent puncture (Figs 37-39) as is 
usual in the Rhadalinae. This character only occurs in Eucymbolus and Rhadalus except for Anthriboclerus 
and Hemipleurus which have some sparse, erect elytral setae emerging from tuberculate punctures. 

The type of Rhadalus testaceus LeConte differs from the species assigned to Cymbolus in its narrower, 
elliptical pronotum, without trace of angles, it denser, tuberculate pronotal punctures (without raised 
shiny, impunctate patches), the longer antennae with more elongate apical segments and its more 
elongate, less setose, tarsi. The epipleura are of a more even width throughout their length; in Cymbolus 
they are much broader at the base than at the level of the hind coxae. These characters are probably only of 
specific significance. 

Crowson (1964) associated Rhadalus LeConte with the other genera of his Aplocneminae on the basis of 
a female specimen from California. The genus was originally placed in the family Melyridae, tribe 
Rhadalini, by LeConte, and was later raised to family rank by Pic (1926), together with the genus 
Dasyrhadus Fall. The latter genus is a dasytine, according to Crowson (1964), and certainly the example of 
Dasyrhadus longior Fall in the BMNH does not belong to the Rhadalinae. Both Hatch (1962) and 
Blackwelder (1975) place Dasyrhadus in the Rhadalinae, which they treat as a subfamily of the Melyridae. 

Crowson also transferred Cymbolus [= Rhadalus] rufopiceus Gorham to the Rhadalinae from the 
Melyrinae. 

Rhadalus is closely related to Eucymbolus (p. 145). 



REVIEW OF THE RHADALINAE 145 

Checklist of species 

castaneus (Gorham), 1886: 324 (from Cymbolus). Mexico. Comb. n. 

elongat us (Champion), 1913: 128 (from Cymbolus). Mexico. Comb. n. 

lecontei Casey, 1895: 605. U.S.A. 

punctipennis (Gorham), 1886: 325 (from Cymbolus). Guatemala. Comb. n. 

quadrituberculatus (Champion), 1913: 129 (from Cymbolus). Brazil. Comb. n. 

rufopiceus (Gorham), 1886: 324 (from Cymbolus). Guatemala. Comb. n. 

testaceus LeConte, 1852: 212. U.S.A. 

wolcotti (Hinton), 1934: 21 (from Cymbolus). Mexico. Comb. n. 

EUCYMBOLUS Champion 

Eucymbolus Champion, 1913: 129. Type species: Eucymbolus cyaneus Champion, 1913: 130, by original 
designation. 

Broadly oval, elytra almost as broad as long, shiny. Body-length less than twice width. Pronotum, elytra 
and 1st visible abdominal sternites metallic blue-black, remainder reddish brown. Dorsum with erect or 
semi-erect long golden setae. 

Head subvertical, frons without grooves or ridges, shiny, with sparse simple punctures. Eyes with sparse, 
short setae, strongly emarginate. Antennae not very long, scarcely reaching shoulder, serrate from 4th 
segment, segments about as long as broad except for apical which is elongate. Apical segment of maxillary 
palps hatchet-shaped, with curved sides. Pronotum subelliptical, strongly transverse, more than twice as 
broad as long; sides explanate and weakly crenulate, anterior edge straight, not bordered in middle, 
posterior edge evenly curved, strongly bordered, all angles rounded; punctures fine and simple on disc, 
tuberculate at sides. Scutellum broadly subtriangular, shiny, sparsely punctured and setose. Elytral sides 
smooth, not explanate, with a carina running from just beneath humeral swelling to about one-third of 
elytral length from apex, not parallel with edge, furthest from it beneath shoulder, scarcely prominent, so 
not forming a shelf; punctures near carina large; elytra strongly punctured, with setae emerging from 
centres of punctures; epipleura similar to Rhadalus but rather broader near base. Tarsi similar to Rhadalus, 
segments 1-4 weakly lobed. Anterior tibiae with short, but distinct, external spines. Prosternal process and 
prosternum similar to that of Rhadalus. Length 4-3 mm (width at broadest part 2-7 mm). 

Distribution. Guatemala. 

Material examined 
Holotype (unique) (BMNH). 

Remarks. The single specimen known of this genus is possibly merely an aberrant Rhadalus, from which it 
differs in its colour, its less elongate form, the shiny sparsely punctured scutellum, the puncturation of the 
pronotal disc and the oblique sublateral carina on the elytra. 

Eucymbolus was transferred from the Melyrinae to the Rhadalinae by Crowson (1964). 

Checklist of species 

cyaneus Champion, 1913: 130. Guatemala. 

ANTHRIBOCLERUS Schenkling 

(Figs 25, 27, 29, 49) 

Anthriboclerus Schenkling, 1922: 328. Type species: Anthriboclerus scotti Schenkling, 1922: 328, by 
original designation. 

Superficially clerid-like. Cuticle unicolorous brown or with indistinct lighter patches on elytral depression 
and lateral pronotal margin. Dorsum shiny, with dark brown, sparse, semi-erect setae interspersed with 
sparse, longer, erect setae. On the elytra these erect setae emerge from the centres of large, weakly 
tuberculate punctures. Elytral tubercles densely setose. Apical antennal segments darker than rest. Tarsi 
yellowish brown. 

Head very large and broad, convex and shiny with a faint diagonal ridge on each side of frons from base 
of antenna to eye; sparsely and minutely punctured (Fig. 25). Eyes very prominent, setose, weakly 
emarginate. Antennae serrate from 4th segment, very long but segments 4-10 not elongate. Maxillary 
palps with apical segment broadly triangular. Pronotum very large and long, about half elytral length, very 



146 E. R. PEACOCK 

strongly convex, shiny, outline from above quadrate, sides markedly explanate, smooth, subparallel; with 
strong posterior border; all angles rounded; punctures minute, simple, very sparse (Fig. 27). Scutellum 
smooth, shiny, unpunctured, shield-shaped. Elytra (Fig. 29) with very prominent humeral swellings and an 
even more prominent setose tubercle (with black setae) on each, near base midway between suture and 
humeral swelling; posterior to these tubercles is a transverse depression and just posterior to this 4 setose 
patches (with very dense white setae) (2 on each elytron); sides smooth; cuticle smooth, shiny, with very 
sparse punctures of two distinct sizes, the larger ones weakly tuberculate or rimmed, with an erect seta 
emerging from the centre of each; epipleura visible from side near base, with smooth inner edge deflecting 
dorsad at about level of 2nd visible abdominal sternite. Hind wing as in Fig. 49. Tarsi 4-segmented, 
segments 1-3 lobed, segment 1 very long, about as long as 4, and longer than 2 and 3 together, segment 3 
slightly smaller than 2; segments 1 and 2 with long ventral setae; claws with large free membranous 
appendages. Prosternal process obsolete; anterior edge of prostemum emarginate to receive head, which is 
sub vertical; prosternum long in front of coxae. Length about 2-3 mm. 

Distribution. Seychelles. 

Material examined 

Holotype and 2 paratypes (BMNH). 

Remarks. Anthriboclerus differs from all other genera in the subfamily by its 4, 4, 4 tarsal formula. It is 
allied to Pelecophora from which it differs mainly in its general appearance, its prominent humeral 
swellings, transverse depression and setose tubercles on the elytra, its sparse dorsal puncturation and 
pubescence, the type of puncturation on the pronotum and elytra, the lack of a prosternal process and the 
ability to fold the head down on to the prosternum. 

Although Anthriboclerus was originally described in the Cleridae and was later listed under the 
Corynetinae (Cleridae) in Corporaal's revised catalogue (1950), Crowson (1964) transferred it to the 
Rhadalinae. 

Checklist of species 

scotti Schenkling, 1922: 328. Seychelles. 

PELECOPHORA Dejean 

(Figs 15, 20, 33, 34, 43, 50) 

Pelecophora Dejean, 1821: 115. Type species: Notoxus illigeri Gyllenhal, 1808: 53, by monotypy. [Dejean 

lists a second species under Pelecophora, but it is a nomen nudum.] 
Pelecophorus; Berthold, 1827: 589. Misspelling. 
Diglobicerus Latreille in Cuvier, 1829: 475 - subgenus erected by Latreille for an unnamed species with 

10-segmented antennae. 

Form elongate, usually with bicolorous cuticle or setae, or both. Setae usually of at least two different 
types, suberect, erect, recumbent or scale-like, the erect and suberect ones usually of a dark colour, the 
recumbent ones usually paler. Pronotal setae directed anteriorly towards the head. 

Head very large and broad, not retractable under pronotum; frons usually with strong subparallel 
grooves, running from above antennal base to eye; strongly and densely punctured with simple or 
tuberculate punctures. Eyes comparatively small, oval, glabrous and very prominent. Antennae mod- 
erately elongate, usually reaching back to base of pronotum, serrate from 4th segment (Fig. 43). Apical 
segment of maxillary palp forming an asymmetrical, elongate, almost right-angled triangle (Fig. 33). 
Pronotum broadest anteriorly or in middle, bordered anteriorly, not always posteriorly; anterior angles 
varying from acutely pointed or right-angled to obtuse-angled or rounded, posterior angles either well 
defined or rounded; disc shiny, punctures simple or, occasionally, weakly tuberculate; punctures tubercu- 
late at sides, surface sometimes rugose near posterior angles (Fig. 20). Scutellum with sides subparallel, 
apex broadly obtuse to semicircular, surface usually with small, dense punctures and scale-like or 
recumbent hair-like pubescence obscuring cuticle. Elytra with sides smooth, weakly explanate; punctures 
large, with long suberect setae, sometimes with patches of very small punctures with scale- or hair-like 
adpressed setae; epipleura broad, visible from side, deflecting dorsad, inner edge becoming obsolete at 
about level of 1st or 2nd visible abdominal seternite. Hind wing as in Fig. 50. Tarsi with segments 1-4 
weakly lobed, segment 5 almost as long as remainder together, much broader at apex; claws with large, free 
membranous appendages. Prosternal process well developed, narrow or stout, sometimes broadening 
posteriorly (Fig. 34), usually on a lower plane than prosternum (about level with ventral side of coxa) and 
completely dividing front coxae. Prosternum very long. Length 3-6 mm. 



REVIEW OF THE RHADALINAE 147 

Distribution. Reunion, Mauritius, E. Africa. [Vinson (1957) expresses doubt as to whether the two E. 
African species (atricolor and jeanneli) described and provisionally placed in Pelecophora by Pic, really 
belong there.] 

Material examined 

13 species, including the type species. 

Remarks. This genus may be recognized by its very large head, entire, prominent eyes, bicolorous cuticle 
or setae, large prosternal process and the tuberculate punctures near the sides of the pronotum. It differs 
from Malthacodes mainly in its large unretractable head, pronotal puncturation and complete prosternal 
process. Its relationship to Anthriboclerus is discussed under that genus. 

Crowson (1964) transferred Pelecophora interrupta Alluaud from the Dasytinae to the Rhadalinae. 

Checklist of species 

albonotata Pic, 1935a: 12. Mauritius. 
albovillosa Vinson, 1946: 259. Mauritius. 
angustataPic, 1935: 116. Mauritius. 
antelmei Alluaud, 1898: 100. Mauritius, Reunion. 

borbonica Vinson, 1967: 330. 
atricolor Tic, 1919: 4. East Africa. 
concinna Vinson, 1946: 260. Mauritius. 
darutyi Pic, 1932: 42. Mauritius. 
decorata Pic, 1911: 151. Reunion. 
emmerezi Pic, 1932: 42. Mauritius. 
hamoni Vinson, 1953: 144. Reunion. 

///jgerj(Gyllenhal), 1808: 53. ?Reunion, Mauritius. (Notoxus) 
obscuricollis Pic, 1932: 43. 
/7/igerii//igerj (Gyllenhal), 1808: 53. ? Reunion, Mauritius. 
Hligeri cariei Tic, 1932: 42; Vinson, 1957: 5. Mauritius. 
illigeri barklyi Vinson, 1957: 5. Round I., Mauritius. 
jeanneliPic, 1919: 3. East Africa. 
marginalis Fairmaire, 1880: 293. Reunion, Mauritius. 
obliquata Alluaud, 1898: 101, Vinson, 1946: 264. 
charmoyi Alluaud, 1898: 102. 
nigrolineata Guerin, 1834: 51. Mauritius. 
pikei Vinson, 1957: 2. Round I., Mauritius. 
subglabra Alluaud, 1898: 99. Mauritius. 
multisignata Pic, 1935a: 12. 
subglabra subglabra Alluaud, 1898: 99. Mauritius. 
subglabra vinsoniPic, 1935a: 12; Vinson, 1946: 269. 
v/tfafa Laporte de Castelnau, 1840: 283. Reunion, Mauritius. 
trimaculata Pic, 1932: 43. 
vittata vittata Laporte de Castelnau, 1840: 283. Reunion. 
vittata interrupta Alluaud, 1898: 101; Vinson, 1958: 119. Mauritius. 

HEMIPLEURUS gen. n. 

(Figs 9, 26, 28, 30, 37, 44, 47, 51) 

Type species: Hemipleurus floriger sp. n. 

Cuticle unicolorous dark brown to black. Head, pronotum and elytra clothed with sparse, semi-erect setae 
interspersed with longer, erect, very stout tuberculate setae, with some recumbent, sparse, whitish setae 
on transverse elytral depression. Ventrally very shiny and sparsely setose. 

Head small, fairly flat, without diagonal lines, retractable against prosternum, anterior edge of which is 
strongly posterior to anterior margin of pronotum; punctures small, rimmed but scarcely visible amongst 
strong microsculpture. Eyes more or less oval, sparsely setose. Antennae moderately elongate, serrate 
from 4th segment (Fig. 44). Apical segment of maxillary palp forming a right-angled triangle. Pronotum 
transverse, all angles broadly rounded, posterior border distinct, sides weakly crenulate, each crenulation 
bearing a long stout seta; punctures rimmed but scarcely discernable. Scutellum broadly semicircular, 
scarcely punctate, more or less glabrous. Elytra more than twice as long as pronotum, shiny, strongly 



148 E. R. PEACOCK 

punctured; with a distinct transverse depression at about one-third of elytral length from base; humeral 
swellings unusually prominent (Fig. 30); epipleura very broad at base and visible from side, shiny, almost 
glabrous, very short, starting to deflect dorsad and becoming obsolete anterior to level of hind coxae (Fig. 
9). Hind wing with radial cell obsolete and anal cell incomplete (Fig. 51). Tarsi with segments 1-4 quite 
strongly lobed, segment 4 smaller than 3; segment 5 broad at apex; claws with large, free membranous 
appendages (Fig. 47). Prosternal process not extending more than half way between front coxae. Length 
2-0-2-3 mm. 

Distribution. Borneo: Sarawak [Mulu]. 

Material examined 

Holotype and paratypes (BMNH). 

Remarks. This genus differs from all other known members of the Rhadalinae except Kubanius by its very 
short inwardly deflecting epipleura. From this genus it is easily separated by the form of tarsal segments 
1-4, which are broad and strongly lobed (Fig. 47) instead of finely elongate (Fig. 10). It resembles 
Anthriboclerus in its peculiarly depressed elytra with sparse erect elytral setae emerging from tuberculate 
punctures, and Malthacodes in its small, deflexed head. From the last genus it differs in not having diagonal 
lines (carinae or grooves) on the head. 

Etymology and gender. Hemipleurus is derived from the Greek 'hemi', meaning 'half and 'pleuron' = 
'pleurus', meaning 'side', referring to the epipleura which are approximately half the body length. The 
name is masculine. 

Checklist of species 
florigersp. n. Borneo. 



Hemipleurus fforiger sp. n. 

Sides of prosternum, epipleura and posterior part of metasternum very shiny, smooth and almost glabrous; 
femora and visible abdominal sternites also very shiny, with sparse semi-recumbent golden setae, with 
some longer dark erect setae near abdominal apex. Tibiae with faint longitudinal microsculpture; sides of 
mesosternum, metasternum and metepisternum with very strong reticulate microsculpture. Colour dark 
brown to black except for basal 3 segments of antennae which are yellowish brown. 

Head transverse, mat due to strong microsculpture, punctures scarcely discernible (Fig. 26); setae 
sparse, very long, black, erect, stout, interspersed with shorter light brown fine semi-erect setae. Eyes 
moderately prominent. Antennae sparsely setose especially near base, weakly serrate from 4th segment, 
segments 4-10 only slightly longer than broad, segments 2-3 narrower, more elongate, subequal in length 
to segments 4-10, segments 1 and 11 longer than others; segments 1-3 light yellowish brown, becoming 
dark brown to black from segment 4 onwards. Labial palps large, apical segment cup-shaped. Pronotum 
mat, due to strong microsculpture (Fig. 28), sides with small setose tubercles, giving a crenulate 
appearance; sculpture unusual: a ring of about 7 small circles around each puncture, these thus resembling 
flowers; setae similar to those on head. Scutellum broad, shiny, semicircular, with fine reticulate 
microsculpture. Elytra with sides smoothly rounded except for a few small tubercles near base, giving 
crenulate appearance; humeral swellings very prominent, with transverse depression posterior to them; 
punctures large, simple, setae similar to those on head and pronotum except for patch of sparse white, 
recumbent setae on elytral depression, which forms a transverse band; clusters of very short erect setae 
emerge from punctures adjacent to most recumbent setae (Fig. 37). 

Material examined 

Holotype $, Sarawak: 4th Division Gn. Mulu NP, v-viii. 1978, on logs nr Base Camp, 50-100 m 

(Hammond & Marshall) (BMNH; B.M. No. 1978-49). 

Paratypes. Sarawak: 5 $ , 4th Division Gn. Mulu NP, v-viii. 1978. 2- on logs nr Base Camp, 50-100 m; 1 
- general sweeping , nr camp 4 , c. 1 800 m ; 1 - nr camp 5 ; 1 - general sweeping , site L 2386 m , upper montane 
forest (Hammond & Marshall) (BMNH; B.M. no. 1978-49). 

Remarks. An unusual feature of this species, not seen on any other species in the subfamily, is the presence 
of clusters of short erect setae adjacent to the recumbent setae on the elytral depression (Fig. 37). These 
could be connected with pheromone glands. 

Etymology, floriger = bearing flowers, referring to the strange pronotal sculpture. 



REVIEW OF THE RHADALINAE 149 

MALTHACODES Waterhouse 

(Figs 8, 35, 41, 42, 46, 52, 53, 54) 

Malthacodes Waterhouse, 1876: 116. Type species: Malthacodes pictus Waterhouse, 1876: 116, by 

monotypy. 
Xamerpus Fairmaire, 1886: 41. Type species: Xamerpus vageguttatus Fairmaire, 1886: 41, by monotypy. 

Syn. n. 
Donaldia Alluaud, 1898: 102. Type species: Donaldia elegans Alluaud, 1898: 103, by monotypy. 

[Synonymized by Vinson, 1958: 123.] 

Generally light-coloured, often with cuticle and/or setae bicolorous, often with erect setae among 
semi-erect or recumbent ones. Sometimes partially metallic. Wing venation varying from weak, without an 
anal cell, to strong, with anal cell distinct (Figs 52-54). 

Head retractable against prosternum, frons with distinct diagonal groove or line on each side, running 
from inner side of antennal base to eye; surface shiny, punctures rimmed. Eyes sparsely setose, varying 
from being very weakly emarginate to entire, fairly prominent. Antennae long, reaching back beyond base 
of pronotum, serrate from 4th segment; varying from being covered with thick, long, black setae, with 
segments 6-10 much broader than long (compressed together and half-moon shaped in the type species 
(Fig. 42)), to sparsely setose and weakly elongate-serrate. Apical segment of maxillary palps triangular. 
Pronotum varying from being very convex in transverse and longitudinal section and at least twice as wide 
as long (in type species) to much less convex and transverse in other species; sides explanate, weakly 
crenulate or smooth, distinctly bordered posteriorly, angles usually rounded, punctures small, rimmed, 
sometimes forming chains, or each (rarely) with an encircling pattern. Scutellum broadly triangular or 
semicircular, shiny, with minute punctures and long fine setae, sometimes so densely punctured and setose 
that cuticle is obscured. Elytra strongly convex, weakly explanate beneath humeral swelling, lateral margin 
sometimes crenulate; punctures strong, becoming larger towards sides, almost forming a stria; epipleura 
broad, shiny, deflecting dorsad and becoming obsolete at level of about 2nd or 3rd abdominal sternite, 
concave and sloping inwards from base so not visible from the side in the broader species, but more 
horizontal and just visible from the side in the more elongate species. Hind wing venation usually weak, 
without anal cell, but occasionally strong with anal cell distinct (Figs 52-54). Tarsi with segments 1-4 
weakly lobed (4 slightly smaller), apical segment broader than others; tarsi small, delicate, usually pale 
yellow or brown and segments 1-3 with very long setae beneath, segmentation often indistinct; [in the 2 
Indian species, M. indicus (Champion) and M. moestus (Gorham), the tarsi are robust and more elongate, 
less strongly setose beneath, with the segments very distinct, the 4th not as small comparatively;] claws with 
large free membranous appendages. Prosternal process small and narrow but extending between front 
coxae and fitting into notch in mesosternum. Length 1-9-4-3 mm. 

Distribution. Madagascar, Seychelles, Aldabra, Mauritius, Reunion, Rodriguez I., Africa, India, Sri 
Lanka. 

Material examined 

25 species, including syntypes of type species {pictus Waterhouse (BMNH), vageguttatus (Fairmaire) 

(MNHN) and elegans (Alluaud) (BMNH)) and some unnamed species from Aldabra. 

Remarks. This genus contains species that are very variable in appearance and which form distinct 
species-groups. I believe that the species formerly assigned to Donaldia are only one such group. The 
differences between these and the type species of Xamerpus are no greater than the differences between 
some of the other species of Xamerpus. For example, X. maindroni (Pic) and X. cioides Champion differ 
markedly from the type species in possessing quite different antennae, a much longer prosternum, a less 
explanate pronotum, more nearly horizontal (less sloping) epipleura as well as a more elongate form. 
Aplocnemus indicus Champion and A. moestus Gorham fall into this group and are here transferred to 
Malthacodes. Apart from the above mentioned characters, these two species have larger, much more 
elongate tarsi than the type species and may prove to merit separate generic status. Vinson (1958) 
synonymized Donaldia with Malthacodes and the only differences found between these two genera is in the 
wing venation, which is strong in the two species originally assigned to Malthacodes, with a distinct anal cell 
(Fig. 52) and weaker in those assigned to Donaldia, with no anal cell (Fig. 54). Most species assigned to 
Xamerpus that I have examined (including the type species (Fig. 53)) have the weak wing venation with no 
anal cell, but one African species has strong venation with an anal cell so, as the other generic characters 
are constant, I am treating this as a variation. The only constant external difference between Malthacodes, 
Donaldia and Xamerpus is that the two former genera have a densely punctured, densely setose scutellum 
and the species of Xamerpus that I have seen have a shiny sparsely setose scutellum. I do not think that this 



150 E. R. PEACOCK 

difference alone is enough to separate the two groups so Xamerpus Fairmaire is here synonymized with 
Malthacodes Waterhouse. 

Malthacodes vageguttatus {Xamerpus vageguttatus) has very similar characters to Malthacodes pictus 
(types compared); with the same general broad, rounded, very convex form, and the head tucked under 
the pronotum. The main differences are the erect and semi-erect dorsal setae, the unusual antennae, the 
crenulate lateral margins of the pronotum and the fainter wing venation with no anal cell in M. 
vageguttatus. 

Crowson (1964) transferred Malthacodes pictus Waterhouse, Donaldia ( = Xamerpus) maindroni Pic 
and X. maculatipennis Pic to the Rhadalinae from the Dasytinae. 

Checklist of species 

alluaudi r (Pic) , 1903: 145 (from Xamerpus). Madagascar. Comb. n. 
ambrensis (Pic), 1931^(445): 101 horstexte (from Xamerpus) . Madagascar. Comb. n. 
bequaerti (Pic) , 1954a: 211 (from Xamerpus) . Belgian Congo. Comb. n. 
bourgeoisii (Fairmaire) , 1898: 476 (from Xamerpus) . Madagascar. Comb. n. 
brevis (Pic), 1931^(445): 101 horstexte (from Xamerpus). Madagascar. Comb. n. 
brunneus (Pic) , 1903: 144 (from Xamerpus). Madagascar. Comb. n. 
cinereovariegatus Blair, 1935: 272. Rodriguez I. 
cioides Champion, 1923: 302 (from Xamerpus) . Seychelles. Comb. n. 
conradsi (Pic), 1939: 169 (from Xamerpus). Tanganyika. Comb. n. 
disconotatus (Pic) , 1931/?: 442 (from Xamerpus) . Madagascar. Comb. n. 
distinctus (Fairmaire), 1901: 180 (from Xamerpus) . Madagascar. Comb. n. 
elegans (Alluaud), 1898: 103 (Donaldia; Vinson, 1958: 123). Mauritius, Reunion. 
e/ega/ise7egajis (Alluaud), 1898: 103. Mauritius. 

elegans bourbonicus (Pic), 1948: 8 (Donaldia, Vinson, 1958: 123; 1967: 330). Reunion. 
elongatus (Pic), 1903: 144 (from Xamerpus) . Madagascar. Comb. n. 
fairmairei (Alluaud), 1898: 103 (from Xamerp us). Madagascar. Comb. n. 
fasciatus (Pic), 1931^(445): 101 hors texte (from Xamerpus) . Madagascar. Comb. n. 
gedyei (Pic), 1938: 300 (from Xamerpus). Kenya. Comb. n. 
indicus (Champion), 1922: 127 (from Aplocnemus). India. Comb. n. 
laterufus (Pic) , 19316: 442 (from Xamerpus) . Madagascar. Comb. n. 
latesuturalis (Pic) , 1958: 205 (from Xamerpus) . Guinea. Comb. n. 
luteofasciatus (Pic) , 1914a: 11 (from Xamerpus) . Madagascar. Comb. n. 
maculatipennis (Pic), 1937: 28 (from Xamerpus), replacement name for 

maculatus Pic, 1926(424): 27 hors texte. E. Africa. Comb. n. 
maindroni (Pic) , 1906: 7 (from Xamerp us). India. (Donaldia) Comb. n. 

pallidithorax (Pic), 1906: 6 (Xamerpus). • 

bimaculatus (Pic), 1906: 8 (Xamerpus). 

unimaculatus (Pic), 1906: 8 (Xamerpus). 

immaculatus (Pic), 1938a: 158 hors texte (Xamerpus). 
martini (Fairmaire), 1898: 476 (Xamerpus). Madagascar. Comb. n. 

maculatus (Pic), 1906: 7 (Xamerpus). 

rufithorax (Pic), 1938a: 158 hors texte (Xamerpus). 
metallicus (Pic) , 1913a: 15 (from Xamerp us). Madagascar. Comb. n. 
minor (Pic), 1932: 46 (Donaldia; Vinson, 1958: 124). Mauritius, Reunion. 

caroli (Pic), 1948: 8 (Donaldia). Mauritius. 
minutus (Pic), 1906a: 11 (Donaldia, Vinson, 1958: 124). Mauritius, Reunion. 

notaticeps (Pic), 1932: 46 (Donaldia). Reunion. 
moestus (Gorham), 1895: 323 (from Aplocnemus) . India. (Dasytes) Comb. n. 
nigriceps (Pic), 19316: 442 (from Xamerpus). Madagascar. Comb. n. 
nigricolor (Pic), 1938a: 158 hors texte (from Xamerpus). Natal. Comb. n. 
nigromaculatus (Pic), 1919: 4 (from Xamerpus). E. Africa. Comb. n. 
obscurus (Pic), 1904a: 28 (from Xamerpus). S. Africa. Comb. n. 
oxylepisiformis (Pic) , 1931a: 107 (from Xamerpus). Madagascar. Comb. n. 
parvus nom. n. for 

minutus Pic, 1931a'(443): 96 hors texte (nee Pic, 1906) (Xamerpus). Madagascar. 
perforatus (Pic), 1917: 5 (from Xamerp us). Sri Lanka. Comb. n. 
perrieri (Fairmaire), 1901: 180 (from Xamerpus). Madagascar. Comb. n. 
pictus Waterhouse, 1876: 116. Rodriguez I. 



REVIEW OF THE RHADALINAE 151 

rubronotatus (Pic) , 1904: 11 (from Xamerpus). Madagascar. Comb. n. 

ruficollis (Pic) , 1931/?: 443 (from Xamerpus). Madagascar. Comb. n. 

sicardi (Pic), 1931c: 447 (from Xamerpus). Madagascar. Comb. n. 

sinuat us (Pic), 1953a: 253. (Donaldia; Vinson, 1958: 123). Madagascar. 

subapicalis (Pic) , 1931d(445): 101 horstexte (from Xamerpus). Madagascar. Comb. n. 

subdepressus (Pic) , 1939: 169 (from Xamerpus). Comb. n. Madagascar. 

subfasciatus (Pic) , 1931a: 108 (from Xamerpus). Madagascar. Comb. n. 

suturalis (Pic), 1931^(445): 101 horstexte (Donaldia, Vinson, 1958: 123). Madagascar. 

trinotatus (Pic), 1938a: 158 hors texte (from Xamerpus). 'B. E. Africa.' Comb. n. 

vageguttatus (Fairmaire), 1886: 41 (from Xamerpus). Madagascar. Comb. n. 

variabilis (Pic), 1931^(445): 101 horstexte (Donaldia: Vinson, 1958: 123). Madagascar. 

interrupta (Pic), 1931^/(445): 101 horstexte (Donaldia). 
vicinus (Pic) , 1931^/(443): 96 hors texte (from Xamerpus). Madagascar. Comb. n. 

INDIODASYTES Pic 

(Figs 17, 24, 45, 55) 

Indiodasytes Pic, 1916: 14. Type species: Dasytes (Indiodasytes) madurensis Pic, by monotypy. Here 
transferred from the Dasytinae (Pic, 1937: 29) to the Rhadalinae. 

Elongate, head and pronotum black, elytra dark brown. Setae paler, either recumbent or semi-recumbent 
and sparsely interspersed with much longer, erect setae, these sometimes occurring only along lateral 
elytral and pronotal margins; pronotal setae point towards centre of disc. 

Head broader than long, subvertical, mat; frons with strong diagonal ridges at inner side of antennal 
insertions; with confused punctures. Eyes prominent, oval, almost glabrous. Antennae serrate from 4th 
segment, these segments not elongate; reaching beyond base of pronotum. Apical segment of maxillary 
palps triangular. Pronotum transverse, sides narrowly explanate and weakly crenulate, bordered post- 
eriorly, strongly punctured with large, rimmed/tuberculate punctures except for a distinct unpunctured 
shiny longitudinal area in mid line at base; intervals between punctures sometimes raised and shiny; angles 
rounded (Fig. 24). Scutellum shield-shaped, mat or shiny, with minute punctures. Elytra with sides weakly 
explanate and crenulate only near base , sinuate when viewed laterally ; epipleura deflecting dorsad at about 
level of 2nd visible abdominal sternite, visible from side near base (Fig. 17). Hind wing of undescribed 
species from Sri Lanka, as in Fig. 55. Tarsi quite stout; claws with large free membranous appendages (Fig. 
45). Prosternal process narrow, but reaching mesosternum between front coxae. Length 2-0-3-7 mm. 

Distribution. India, Sri Lanka. 

Material examined 

Holotype of type species (MNHN) and 2 undescribed species from Sri Lanka (BMNH). /. pubicornis 
Wittmer from China, which was doubtfully placed in this genus by Wittmer (1940), does not belong to it 
and is not a rhadaline. 

Remarks. This genus strongly resembles Aplocnemus in general appearance but differs in its rimmed 
pronotal puncturation, its diagonal ridges on the head and from most species by its strongly triangular 
apical segment of the maxillary palps. It also resembles Malthacodes but differs from most species in its 
proportionally larger, less strongly setose tarsi and its stronger pronotal puncturation. It differs from all 
other genera in the Rhadalinae, that I have examined, by the lack of a dorsal appendage to the median lobe 
of the aedeagus. However, its other characters, e.g. first two visible abdominal sternites fused, triangular 
apical segment to the maxillary palps, free membranous appendages to the claws etc. , show that it belongs 
to this group. Indiodasytes was listed in the Dasytinae by Pic (1937). It is here transferred to the 
Rhadalinae. 

Checklist of species 

madurensis (Pic), 1916: 14. (Dasytes.) India. 

APLOCNEMUS Stephens 

(Figs 14, 19,31,32,56) 

Elicopis Stephens, 1829: 136. Type species: Crioceris impressa Marsham, 1802: 226, designated by 
Stephens, 1830: 316 for the replacement name Aplocnemus. [See also Note below.] 



152 E. R. PEACOCK 

Aplocnemus Stephens, 1830: 316. Type species: Crioceris impressa Marsham, 1802: 226, by original 

designation. [Replacement name for Elicopis - see Note below.] 
Haplocnemus; Agassiz, 1846[7]: 29, 172. [Unjustified emendation of Aplocnemus Stephens.] 
Helicopis; Agassiz, 1846[7]: 29, 175. [Unjustified emendation of Elicopis Stephens and junior homonym of 

Helicopis F. 1807 (Lepidoptera: Riodinidae).] 
Diplambe Schilsky, 1894: 234 (s.g.). Type species: Dasytes montivagus Rosenhauer, 1856: 156, by 

PRESENT DESIGNATION. 
Holcopleura Schilsky, 1894: 234 (s.g.). Type species: Haplocnemus (Holcopleura) reitteri Schilsky, 1894: 

234, by monotypy. 
Ischnopalpus Schilsky, 1894: 235 (s.g.). Type species: Haplocnemus (Ischnopalpus) subcostatus Schilsky, 

1894: 235, by subsequent designation (Schilsky, 18946: no. 62). 
Pseudaphyctus Pic, 1896: 47 (s.g.). Type species: Haplocnemus (Pseudaphyctus) toumieriYic, 1896: 47 [= 

Haplocnemus tumidus Kiesenwetter, 1863: 650], by monotypy. 

Note. Thomson (1859: 109) designated Lagria nigricornis Fabricius (1792: 81) as type species of 
Aplocnemus. This was not only unnecessary but invalid, because it was not included in the original 
description of Aplocnemus . 

The present genus was first recognized by Stephens under the name Elicopis (1829: 136), but later, he 
thought that the name was 'too closely allied to Hellicopis" [= Helicopis] F. (1807), and he changed it to 
Aplocnemus (1830: 316). Since the latter name has been in use for over 150 years, I suggest that it is 
conserved on the grounds of long usage (1985, International Code of Zoological Nomenclature, Article 
23(b)). In the interest of stability, a case should be made to the International Commission. 

Unicolorous brown to black (mostly black), often with a metallic bluish or greenish tinge, legs and base of 
antennae often lighter. Dorsum usually covered with erect or semi-erect long fine setae, legs and 
abdominal sternites with similar but more recumbent setae; almost invariably with a few very long, erect 
setae on tibiae among the recumbent ones; setae usually black but sometimes grey, golden or white. Tibiae 
often with spines. Elytral setae sometimes alternately erect and semi-erect. 

Head moderately small, shiny or mat with strong simple punctures; frons usually with two short parallel 
depressions at inner side of antennal insertions. Eyes prominent, sparsely setose, oval or weakly 
emarginate. Antennae serrate or pectinate from 3rd or 4th segment, apical segment elongate-oval, length 
varying from barely reaching base of pronotum to reaching well beyond it. Apical segment of maxillary 
palps varying from spindle-shaped (rarely) (Fig. 32) to hatchet-shaped (Fig. 31). Pronotum transverse; 
sides narrowly explanate, finely crenulate or smooth, usually bordered anteriorly and posteriorly; base 
sinuate; disc finely and sparsely or strongly and densely punctured with simple punctures (Fig. 19), only 
rarely punctures weakly rimmed on disc or at sides. Scutellum shiny, quite strongly punctured, broadly 
triangular or semicircular. Elytra with sides weakly explanate near base, crenulate or smooth (in subgenus 
Diplambe Schilsky the elytra are double-sided in basal half, i.e. a carina runs subparallel to elytral edge, as 
also in Rhadalus); epipleura deflecting dorsad at about level of 2nd, 3rd or 4th visible abdominal sternite, 
often visible from side near base. Hind wing venation variable, but usually distinct (Fig. 56). Tarsi 
moderately elongate, segments 1-4 weakly lobed; claws with large free membranous appendages which 
sometimes appear to be sclerotized along side nearest claw (Fig. 14). Legs moderately slender. Prosternal 
process apparently short, not separating front coxae, usually with fine process which extends for length of 
coxa but is hidden from below by contiguous coxae. Length 2-9-6-9 mm. 

Distribution. Palaearctic (including N. Africa), Asia Minor, W. Africa (Ivory Coast, Angola), Philippine 
Is (1 species). 

Material examined 

About 100 species (including syntypes of type species (BMNH)). 

Remarks. This genus is very large and variable, mostly with the general appearance of being black and 
hairy. Its most constant characters are the tarsi and claws, head and maxillary palp shape and the (almost 
constant) long suberect setae. Most species are readily recognized by their dark coloration, elongate form 
and hairy appearance. They are usually dark brown or black, sometimes metallic. One species from the 
Canary Is differs from the majority in its almost recumbent dorsal setae. Other Canarian species tend to be 
of a light yellowish brown colour wholly or in part. Most species resemble Trichoceble in general 
appearance but may be easily distinguished by the presence of membranous appendages and the lack of 
chitinised teeth on the claws, as well as by the less strongly emarginate eyes. They may be distinguished 
from all other genera except Semijulistus and Microjulistus by the only moderately securiform apical 
segment of the maxillary palps (spindle-shaped in some species). Most other genera have distinctly 



REVIEW OF THE RHADALINAE 153 

triangular apical segments. They differ from Malthacodes in their simple (non-rimmed) pronotal punctures 
and the lack of diagonal ridges on the head. (The Indian species, Aplocnemus indicus Champion and 
moestus Gorham are here transferred to Malthacodes. They differ from Aplocnemus in their weaker wing 
venation, triangular apical segment of the maxillary palps, diagonal lines on head, pronotal puncturation 
and the form of the male genitalia). 

Crowson (1964) verified that Aplocnemus palaestinus Baudi, A. (Diplambe) abietum Kiesenwetter and 
A. (Ischnopalpus) subcostatus Schilsky belong in the Rhadalinae. They were previously placed in the 
Dasytinae (Pic, 1937). 

Checklist of species 

abietum Kiesenwetter, 1859: 171. Mediterranean Region. 
acutangulus Schilsky, 1897a: no. 92. Algeria, Italy. 
adanensis Pic, 1908a: 50. Turkey. 
aerosus Schilsky, 1897a: no. 97. Spain. 
aestivus Kiesenwetter, 1863: 654. Europe. 

subviolaceus Pic, 1922: 27. Austria, 
a/er Schilsky, 18976: no. 33. Algeria. 
akbesianus Pic, 1896: 48. Syria. 
albipilis Kiesenwetter, 1863: 651. Spain. 
algiricus Schilsky, 18946: no. 73. Algeria. 
alluaudiVic, 1900: 165. Tunisia. 
alpestris Kiesenwetter, 1861: 385. Europe. 

pectinicornis (Dufour), 1851: 329. (Dasytes) 

tarsalis; Schilsky, 18976: no. 34LL; Lohse, 1977: 178; Majer, 1982: 435. S. Europe. 

theresae Pic, 1914: 57. France. 
alternates Peyerimhoff , 1931 (1932): 1. Morocco. 

cribripennis Pic, 1922: 27. Algeria. 

nigrescens Pic, 1949: 74. 
anatolicus Schilsky, 1903: no. 97. Turkey. 

subelongatus Pic, 1910: 9. 
andalusicus (Rosenhauer), 1856: 158. Spain. (Dasytes) 

laetus Schilsky, 18976: no. 34HH. 
angolan us Wittmer, 1953: 285. Angola. 
atricornis Pic, 1921: 3. Greece. 
aubei Kiesenwetter, 1867: 122. Pyrenees, Spain. 
baborensis Pic, 1908: 200. Algeria. 

baborensis Pic, 1922: 26. Algeria. [Homonym of baborensis Pic 1908.] 
barnevillei Kiesenwetter, 1867: 121. Spain. 
basalis (Kiister), 1849: no. 19. S. Europe. (Dasytes) 

distinctipes Pic, 1908a: 50. Corfu. 
beaupreiYic, 1909: 105. Tunisia. 
berytensis Sahlberg, 1913: 45. Syria. 
bicoloratus Pic, 19356: 15. Morocco. 

theryi Pic, 19356:15. 
biskraensis Schilsky, 1897a: no. 96. S. Algeria. 

biscrensis Schilsky, 18976: no. 34QQ. 

biskrensis Schilsky, 18976: no. 63. 

gridettii Pic, 1928: 103. 
brevis (Rosenhauer), 1856: 161. Spain. (Dasytes) 
brevissimus Pic, 1908a: 50. Greece. 

holtziYic, 1908a: 50. 
breviusculus Schilsky, 18976: nos 31 & 34CC. 
caelatus Brulle, 1832: 151. Greece. 
ca/idus Mulsant & Rey, 1868: 235. S. France, Italy. 
capillicornis Abeille, 1907: XXI. S. France. 
caramanicus Sahlberg, 1913: 46. Caramania. 
caroli Pic, 1941: 1. Morocco. 
castiliensis Schilsky, 18976: nos 41 & 34FF. Spain. 
caucasicus Schilsky, 1897a: no. 89. U.S.S.R.: Caucasus. 



154 E. R. PEACOCK 

chalconatus (Germar), 1817: 209. S. Europe. (Dasytes) 

aestivus Kiesenwetter, 1863: 654 (partim); Majer, 1982: 442. Europe. 

pinicola Kiesenwetter, 1863: 653 (partim females) nee pinicola: Lohse, 1977; Majer, 1982: 442. 
S. Europe. 
chlorosoma (Lucas), 1849: 199. N. Africa, Italy, Sardinia. (Dasytes) 

cuprea (Lucas), 1849: 199. 
cobosi Pic, 1953: 144. Spain. 

coeruleatus (Rosenhauer), 1856: 160. Spain. (Dasytes) 
consobrinus (Rosenhauer) , 1856: 157. Spain. (Dasytes) 
corcyrkus Miller, 1866: 818. Corfu, S. Italy. 
crenicollis Kiesenwetter, 1863: 654. Italy, Sardinia. 

duplicatus Kiesenwetter, 1871: 85. 
cribrarius (Brulle), 1832: 151. Greece. (Dasytes) 
cribrkollis Mulsant & Rey, 1868: 234. Corsica, Sardinia. 
cribripennis Tic, 1921: 3. Greece. 
cribrosus Schilsky, 18976: nos. 44 & 34GG. Algeria. 

festai Pic, 1925:2. 

robustior Pic, 1928: 103. 
croceicornis Kiesenwetter, 1863: 649. U.S.S.R.: Sarepta. 
cupreatus Schilsky, 1897a: no. 88. U.S.S.R.: Caucasus. 
curticornis Pic, 1910: 9. Morocco. 
curtipennis Pic, 1908: 200. Algeria. 
curtus Pic, 1921: 3. Algeria. 
cylindricus Kiesenwetter, 1863: 651. S. Europe. 
cyrenaicus Pic, 1955: 132. Libya. 
delagrangei Pic, 1902: 32. Syria. 
dentatus Schilsky, 1897a: no. 100. Algeria. 
depressicollis Schilsky, 18976: nos 52 & 34LL. Kurdistan. 
desertorum Pic, 1896: 48. Algeria. 

rufofemoratus Pic, 1908a: 49. 
diaphanus Schilsky, 1897a: no. 94. Algeria, 
di/fici/is Holdhaus, 1923: 137. Italy. 
esca/erai Pic, 1908a: 90. Spain. 
escalerai escalerai Pic, 1908a: 90. 
escalerai co/asi Pic, 1954: 97. 
eumerus Mulsant & Rey, 1868: 194. France. 

marchali Pic, 1914: 57. 
fauconneti Tic, 1914: 57. C. France. 
ttavkornis Schilsky, 1987a: no. 87. U.S.S.R.: Caucasus. 

areschanus Pic, 1908a: 81. 
fortepunctatus Pic, 19356: 15. Morocco. 
geniculat us Schilsky, 1903: no. 98. Turkey. 
gestroi Schilsky, 18976: nos 58 & 34NN. Algeria, Tunisia. 
gracilicornis Schilsky, 19876: nos 36 & 34DD. Spain. 
grancanariensis Lindberg, 1953: 7. Canary Is. 
griseopubescens Tic, 1899: 259. Palestine (sic). 
hebraicus Schilsky, 1906: no. 13. Palestine (sic). 
heydeni Schilsky, 18946: no. 81. Algeria, Turkey. 
hickeri Pic, 1935c: 256. Turkey. 
hierichunticus Sahlberg, 1913: 42. Palestine (sic). 
hispankus Pic, 1953: 143. Spain. 
imperforatus Tic, 1908a: 49. Tunisia. 
impressipennis Tic, 1921: 5. Italy. 
impressus (Marsham), 1802: 226. N. & C. Europe. (Crioceris) 

pini (Redtenbacher), 1849: 335. (Dasytes) 

serratus (Redtenbacher), 1849: 335. (Dasytes) 
incognitas (Faldermann), 1836: 204. U.S.S.R.: Caucasus. (Dasytes) 
integer Baudi, 1873: 302. Italy. 
jejunus Kiesenwetter, 1863: 652. S. Europe, N. Africa. 



REVIEW OF THE RHADALINAE 155 



kaszabi Ma jer, 1982: 431. C. Europe. 

kiesenwetteri Schilsky, 1897b: nos 50 & 34KK. Greece, S. Europe. 

korbi Schilsky, 18976: nos 49 & 34HH. Spain. 

diversus Schilsky, 18976: nos 49 & 34HH. 
koziorowiczi Desbrochers des Loges, 1870: 122. Corsica, Italy. 

xanthopits Kiesenwetter, 1871: 85. 

rufomarginatus Schilsky, 1894/?: no. 85. 
krugeri Pic, 1929: 94. Cyrenaica (sic). 
kubanensis Pic, 1909: 99. U.S.S.R.: Caucasus. 
lacoi Majer, 1985: 35. C. Europe. 
lateralis Schilsky, 18946: no. 71. U.S.S.R.: Caucasus. 
latior Pic, 1908a: 50. Greece. 
latipennis Pic, 1911: 146. Syria. 

rechmayanus Pic, 1911: 169. Lebanon. 
libankus Pic, 1901: 9. Syria. 

laetipes Abeille, 1907: XXII. 
limbipennis Kiesenwetter, 1865: 383. Spain. 
longicornis Kocher, 1962: 196. Morocco. 
longulus Schilsky, 1897a: no. 91. N. Africa, Italy. 

brunnescens Pic, 1922: 27. 
macedonicus Pic, 1922: 17. Macedonia. 
mancinii Tic, 1927: 14. Capri. 

cap raianus Pic, 1931: 159. 
marginicollis Schilsky, 18976: nos 39 & 34EE. Algeria. 
tnaurus Schilsky, 1895: no. 18. Algeria. 
melitensis Schilsky, 18976: nos 60 & 3400. Malta. 
mira bilis Schilsky, 1897a: no. 99. Algeria. 
mohamed Chobaut, 1898: 78. S. Algeria. 

montivagus (Rosenhauer) , 1856: 156. Spain, Sardinia. (Dasytes) 
morio (Schonherr), 1817: 12. Morocco. (Dasytes) 
nevadensis Tic, 1908a: 90. Spain. 
nigricornis (Fabricius) , 1792: 81. N. & C. Europe. (Lagria) 

femoralis Illiger, 1807: 302. 

punctatus (Germar), 1824: 77. (Dasytes) 

chalybeus (Germar), 1824: 78. (Dasytes) 

fuscitibia Mulsant & Rey, 1868: 247. 

variolatus Costa, 1882: 35. 

'femoralis' Schilsky, 1897 - incorrect subsequent spelling oifemoratus. 

femoratus Schilsky, 18976: nos 46 & 34FF; Majer, 1982: 430. Hungary. 

viertli Schilsky, 18976: nos 46 & 34FF; Majer, 1982: 430. 

rufolateralis Pic, 1914: 57. 

testaceofemoralis Pic, 1917: 22. 
nigripes Mulsant & Rey, 1868: 223. Algeria. 
palaestin us B audi, 1873a: 319 (Note 1). Israel, Syria. 
parumpunctatus Schilsky, 18976: nos 53 & 34LL. U.S.S.R. 
pectinatus (Kiister) , 1849: no. 18. Sardinia, Corsica, Capri. (Dasytes) 

erosus Mulsant & Rey, 1868: 233. 
pectinicornis (Lucas), 1849: 200. Algeria. (Dasytes) 
pellucens Kiesenwetter, 1865: 383. Spain. 
perforates Schilsky, 18976: nos 35 & 34CC. Algeria. 
persicus Schilsky, 18976: nos 37 & 34DD. Iran. 
pertusus Kiesenwetter, 1859: 172. Greece, Turkey, Cyprus. 

elongatior Pic, 1902: 32. 

adaliensis Pic, 1908a: 50. 
pesruchesi Schilsky, 1897a: no. 90. Algeria. 
peyroni Pic, 1899: 259. Syria. 
philippinus Wittmer, 1941: 226. Luzon. 
ponferradanus Pic, 1913: 105. Spain. 
pristocerus Kiesenwetter, 1859: 170. Greece, Corfu. 



156 E. R. PEACOCK 

pulverulentus (Kiister), 1849: no. 17. Hungary, Turkey, Dalmatia, Italy. (Dasytes) 

obscurus Germar, 1817: 209. 
puncticollis Sahlberg, 1913: 40. Turkey. 
punctiger Schilsky, 1897a: no. 98. U.S.S.R.: Caucasus. 
quercicola Mulsant & Rey, 1868: 235. C. France. 
raffrayi Pic, 1908: 51. Italy. 
ragusae Schilsky, 18946: no. 68. Italy, 
raiiiicorii/s Kiesenwetter, 1863: 651.U.S.S.R.: Sarepta. 
rajmoiidiDeville, 1908: 218. Corsica. 

xanthopus Schilsky, 18976: nos 40 & 34EE. 
reitteri Schilsky, 1894: 234. Turkey. 
rufipes Miller, 1862: 345. Greece, Macedonia. 

taygetanus Pic, 1908a: 50. 

semicaeruleus Pic, 1922: 17. 
rufomarginat us Perris, 1869: 18. N. Africa, Italy, Corsica, Sardinia, S. France. 

marginatus Rottenberg, 1870: 243. Algeria. 

edoughensis Pic, 1897: 41. 
rugicollis Schilsky, 1897a: no. 95. Algeria. 
rugulosus (Rosenhauer), 1856: 159. Spain. (Dasytes) 

subcoeruleus Pic, 1908a: 90. 

mateui Pic, 1954:97. 
russicus Reitter, 1890: 148. U.S.S.R.: Caucasus. 
sanctus Pic, 1902: 32. Palestine (sic). 
sculpturatus Wollaston, 1862: 447. Canary & Madeira Is. 

tenerifensis Pic, 1922: 17. 
serbicus Kiesenwetter, 1863: 653. Yugoslavia, Hungary, U.S.S.R.: Crimea. 

tibiellus Schilsky, 18976: no. 34MM. U.S.S.R. 
serratus (Brulle), 1832: 152. Greece. (Dasytes) 
serrulatus Schilsky, 1906: no. 100. Spain. 
siculus Kiesenwetter, 1863: 654. Italy, Algeria, Tunisia. 

fulvipes Schilsky, 1894: 230. 

flavipes Schilsky, 18976: no. 59. 

obscuripes Schilsky, 18976: no. 59. 
simitis Schilsky, 1906: no. 99. Turkey. 
smyrnensis Pic, 1901: 18. Turkey. 
standi Marcu, 1936: 518. NE. Rumania. 
subcostat us Schilsky, 18946: no. 62. N. Africa. 

bonnaireiYic, 1894a: 112. 

viberti Pic, 1922:27. 
subinteger Tic, 1902a: 60. Mesopotamia. 
suggara Peyerimhoff , 1929: 193. Algeria, Egypt. 
suggara suggara Peyerimhoff , 1929. Algeria. 
suggara ai/ieri/Wittmer, 1938: 169. Egypt. 
syriacus Schilsky, 1894: 235. Syria, Israel. 

caerulescens Schilsky, 18976: no. 34DD. 
tarsalis (Sahlberg), 1822: 113. N & C. Europe. (Dasytes) 

rufitarsis (Sahlberg), 1822: 113. (Dasytes) 

virens (Kiister), 1849: no. 21; Lohse, 1977: 178. (Dasytes) 

pinicola Kiesenwetter, 1863: 653; Lohse, 1977: 178; partim females (nee pinicola: Lohse, 1977): 
Majer, 1982: 442. 

kuesteri Schilsky, 1894a: 331; Lohse, 1977; 178. Germany. 

pandellei Pic: 1918: hors texte 7. 
tarsicola Sahlberg, 1913: 43. Syria. 
testaceipesYxc, 1932a: 27. Morocco. 
thalensisPic, 1930: 1. Tunisia. 
thessalicus Pic, 1908a: 49. Greece. 
tingitanus B audi, 1873: 300. Morocco. 

lixensis Bourgeois, 1911: 158. 

subacuminatus Pic, 1934: 21. 



REVIEW OF THE RHADALINAE 157 

trinacriensisRagusa, 1872: 83. Italy, N. Africa. 
fu&ercii/j/erMotschulsky, 1850: 364. Germany. 
tumidus Kiesen wetter, 1863: 650. Morocco, Spain. 

tournieri Fie, 1896: 47. 
tuniseus?ic, 1908a: 50. Tunisia. 
turcicus Schilsky, 1897/?: nos 30 & 34CC. Turkey. 
uhagoni Schilsky, 1897a: no. 86. Spain. 
venustulus Schilsky, 1897a: no. 93. U.S.S.R.: Caucasus. 
vestitus Wollaston, 1862: 447. Canary Is. 
virens (Suffrian) , 1843: 337. S. & C. Europe. (Dasytes) 
ahenus Kiesenwetter, 1863: 652. 
tibialis Schilsky, 1894a: 331. 

virens virens (Suffrian), 1843: 337. S. & C. Europe. 

virens orientalis Lohse , 1977: 178. W. Germany. 
viridescens Pic, 19356: 15. Morocco. 

KUBANIUS Majer 

(Figs 10, 11,40) 

Kubanius Majer, 1983: 385. Type species: Kubanius elegans Majer, 1983: 387, by original designation. 

Body with a metallic sheen, sparsely punctured; of rather flattened, soft-bodied appearance, with fine, 
elongate legs and tarsi. Body, femora and tibiae sparsely clothed with long, fine suberect setae, tibiae with 
some fine spines among the setae; tarsi with shorter setae and some short spines. Abdominal sternites with 
fine, sparse, recumbent setae. [Of the 2 species seen, one, K. elegans, is unicolorous black, the other 
(undescribed) has a brown to black, metallic, head, pronotum and ventral side with yellowish brown elytra 
and antennae and light yellow legs.] 

Head large, broader than or nearly as broad as anterior part of pronotum; punctures small, sparse, 
rimmed or simple; cuticle with microsculpture or smooth and shiny; short longitudinal groove present, on 
each side of frons at inner side of antennal insertion. Eyes weakly reniform, sparsely setose. Antennae very 
long or of moderate length, not markedly sexually dimorphic, reaching back well beyond base of elytra, 
segments 5-10 large and strongly serrate (Fig. 40). Apical segment of maxillary palps securiform. 
Pronotum small, subquadrate or elliptical, weakly transversely convex, almost flat on disc; sides weakly 
explanate, lateral margins smoothly rounded; anterior angles rounded, posterior angles either distinct or 
obsolete; posterior margin strongly bordered; punctures small, sparse, simple, rimmed or weakly 
tuberculate on disc; cuticle between punctures shiny, with or without microsculpture. Scutellum shiny, 
semicircular, sparsely punctured. Elytra flat on disc, broadening out at sides from shoulder in a weak curve 
and turned down sharply at apices, the widest part being at about one-third of elytral length from apex; 
punctures strong, cuticle transversely rugose; epipleura very short, deflecting dorsad well before level of 
hind coxae. Hind wing venation similar to that of Trichoceble (figured by Majer, 1983). Tarsi (hind) long, 
subequal in length to tibiae, with spiky ventral combs; claws with large blunt teeth and very small, 
inconspicuous membranous appendages (Figs 10, 11)). Tibiae with fine spines. Prosternal process not 
produced between front coxae, which are very strongly projecting and contiguous. (Metasternum with 
very strong median longitudinal groove.) Length 2-0-4-1 mm. 

Distribution. U.S.S.R. (Kazakhstan), N. India. 

Material examined 

1 paratype (BMNH) [the holotype is in the National Museum, Prague and there are more paratypes in 

Majer's collection], and 6 examples of an undescribed species from N. India (BMNH). 

Remarks. Majer used the sharply declining apices of the elytra to separate Kubanius from other genera, 
but as the elytra are rather soft it is possible their curvature may be due to the way the insects dried out after 
killing. However, the genus is a very distinctive member of the Rhadalinae , closely resembling Trichoceble 
in the form of the tarsi and subparallel grooves on the head. The antennae also are very similar to those of 
some species of Trichoceble but the pronotum is much less transverse and convex, the elytra less densely 
punctured, the prosternal process is obsolete and the epipleura much shorter. Hemipleurus gen. n. also has 
short, inwardly deflecting epipleura but is otherwise dissimilar. 

Checklist of species 

elegans Majer, 1983: 387. U.S.S.R. (Kazakhstan). 



158 E. R. PEACOCK 

TRICHOCEBLE Thomson 

(Figs 12, 13,23,39,57) 

Trichoceble Thomson, 1859: 109. Type species: Melyris floralis Olivier, 1790: 11, by original designation. 
Julistus Kiesenwetter, 1859: 174. Type species: Julistus funera Kiesenwetter, 1859: 175, by PRESENT 
DESIGNATION. 

Unicolorous black or dark brown or with legs and antennal base lighter, clothed with fine erect or 
semi-erect unicolorous setae, ranging from golden or grey to black. Ventral pubescence recumbent. Tibiae 
with short setae. 

Head smooth or with an insignificant, short longitudinal groove on each side of frons at inner side of eye 
and antennal insertion; punctures small, sparse, simple. Eyes emarginate, either strongly setose or 
glabrous. Antennae very long and strongly pectinate or serrate from 3rd to 5th segment (some females with 
shorter antennae than males but these still extend well back beyond shoulder). Apical segment of maxillary 
palps longitudinally triangular. Pronotum strongly transverse, convex, subelliptical, with very rounded 
smooth or finely crenulate sides; either all angles rounded or some obtuse; posterior border present; cuticle 
smooth, shiny, sometimes rugose near base, punctures small on disc (Fig. 23), becoming larger towards 
sides, usually simple but occasionally rimmed and becoming weakly tuberculate near sides. Elytra 
gradually broadening from shoulder so that broadest part is about one-third of elytral length from apex; of 
rather smooth appearance, densely punctured, punctures tending to be linked transversely by shallow 
grooves, forming transverse lines; lateral margins very narrowly explanate; humeral swellings moderately 
prominent; epipleura fairly long, weakly deflecting dorsad along their length, so not visible from side, 
evanescing at about level of 2nd abdominal sternite. Hind wing as in Fig. 57. Tarsi (hind) not as long as 
tibiae (cf. Kubanius); claws appendiculate, apices of teeth usually pointed (Figs 12, 13) but occasionally 
blunt. (Legs slender.) Prosternal process projecting about half way between front coxae and sloping to a 
lower plane than prosternum. Length 3-0-7-0 mm. 

Distribution. C, E. and S. Europe, Turkey, Armenia, Kurdistan, Caucasus, Turkestan, Tibet, India, 
China. 

Material examined 

12 species (including type species). 

Remarks. Trichoceble is very closely related to Kubanius (q.v.) and Antinea. It differs from the latter in 
possessing membranous wings, larger reniform eyes and shorter temples. It differs from Aplocnemus in its 
lack of membranous appendages on the claws. Trichoceble cincta Pic from Kurdistan could possibly be a 
different genus as its wing venation is much weaker and no anal cell is present. It also differs in having a 
narrower pronotum, strongly rimmed punctures with a pattern round each one (making them look 
flower-like). Also, it is light brown, except for the black head, which is not the usual coloration in 
Trichoceble. 

Seidlitz (1891) synonymized Trichoceble and Julistus. Although they were both described in the same 
year, Trichoceble was published first. Thomson's book (containing his description of Trichoceble) is 
reviewed in the 'Neuere Literature' of the same part of Berliner Entomologische Zeitschrift, vol. 3 as that in 
which Julistus is described. Crowson (1964) transferred Trichoceble to the Rhadalinae from the Dasytinae, 
on the basis of the examination of T. floralis (Olivier) and Julistus [= T.J oertzeni Schilsky. He also 
verified that the larval characters of T. memnonia (Kiesenwetter) were similar to those of Aplocnemus . 

Checklist of species 

araratica Yablokov-Khnzoryan, 1978: 243. Armenia. 

arbustorum (Kiesenwetter), 1859: 176. Greece. (Julistus) 

cincta?ic, 19296: 1. Kurdistan. 

convexa Pic, 1909: 113. Syria. 

curta (Baudi), 1873: 297. Dalmatia, Turkey. (Julistus) 

elongata Schilsky, 1897a: no. 85. 

floralis (Olivier), 1790: 11. C. & S. Europe, U.S.S.R. (Melyris) 

floricola (Kiesenwetter), 1861: 386. (Julistus) 
/u/vopi7is (Reitter), 1889: 25. U.S.S.R.: Caucasus. (Julistus) 
funera (Kiesenwetter), 1859: 175. SE. Europe, Greece, Italy. (Julistus) 
grandis Schilsky, 1896: nos 90 & 32P. Syria. 
griseohirta (Reitter), 1885: 380. Crete. (Julistus) 
heydeni Schilsky, 1897: 155. China. 



REVIEW OF THE RHADALINAE 159 

immarginata Reitter, 1902: 257. Turkey. 

/aefa Majer, 1986: 303. Mongolia. 

lederi Schilsky, 1896: nos92&32P. U.S.S.R.: Caucasus. 

longicornis (Kiesenwetter), 1863: 647. Syria. (Julistus) 

major Pic, 1922: 17. Greece. 

mediocris Ma jer, 1986: 306. Mongolia. 

memnonia (Kiesenwetter), 1861: 385. C. Europe, Germany, France, U.S.S.R.: Caucasus. (Julistus) 

fulvohirta (Brisout de Barnville), [1862]: 601; Lohse, 1977: 182. (Julistus) 

semirufescens (Pic), 1915: 22. (Julistus) 
nigra Pic, 1921: 4. Greece. 
oculata Schilsky, 1896, nos 98 & 32Q. Greece. 

ocularis Reitter, 1902: 260. 
oertzeni Schilsky, 1896: nos 91 & 32P. Crete. 
pallidipes (Pic) , 1944: 12. China. (Julistus) 
ramicornis Schilsky, 1900: no. 4. Turkestan. 
schatzmayeri Tic, 1909: 113. Macedonia. 
schilskyi Reitter, 1902: 259. U.S.S.R.: Caucasus. 
sparsepunctata Pic, 1921: 4. Turkey. 
subcaerulea Pic, 1921: 4. Greece. 
subcoriacea Reitter, 1902: 260. Greece. 
testaceipes Pic, 1921: 4. Cyprus. 
torretassoi Wittmer, 1935: 254. Rhodes. 
unguicularis Reitter, 1902: 258. Turkey. 
unicolor Pic, 19326: 251. China. 

ANTINEA Peyerimhoff 

Antinea Peyerimhoff, 1929: 191. Type species: Antinea saxicola Peyerimhoff, 1929: 191, by monotypy. 

Unicolorous dark brown. Dorsum covered with fairly short semi-erect setae; humeral swellings obsolete. 
Apterous. 

Head large, nearly as long as broad, temples long, at least twice length of eye when seen from above; 
frons flat or with a weak depression at each side between eyes; shiny with small, simple punctures. Eyes 
small, flat, oval. Antennal segments 3-6 weakly and 7-10 strongly serrate, segment 11 elongate-oval. 
Pronotum strongly transverse, elliptical, all angles rounded, sides rounded, margins crenulate, strongly 
punctured with small simple punctures, without obvious borders. Scutellum subsemicircular, with strong 
puncturation. Elytra with sides curved, narrowly explanate and finely crenulate; humeral swellings 
obsolete; epipleura extending to about level of 2nd visible abdominal sternite. Hind wings absent. Tarsi 
elongate, segments 1-4 weakly lobed, claws fine, with minute teeth. Legs slender with long, stout tibial 
spurs; hind tibiae longer than front and middle tibiae. Prosternal process very short, not produced between 
front coxae. Metasternum unusually short, distance from middle coxa to hind coxa about as long as the 
longitudinal diameter of a hind coxa. Length 4 mm. 

Distribution. N. Africa (Sahara). 

Material examined 
Syntype(BMNH). 

Remarks. Antinea is most closely related to Trichoceble (q.v.). It was listed in the Dasytinae by Pic (1937) 
and is here transferred to the Rhadalinae. 

Checklist of species 

saxicola Peyerimhoff , 1929: 191. N. Africa (Sahara). 

SEMIJUL1STUS Schilsky 

(Figs 1,3, 5, 6, 7, 38, 58) 

Semijulistus Schilsky, 1894: 227 (included in the rhadaline group by Majer, 1983: 387). Type species: 
Dasytes callosus Solsky, 1867: 32, by original designation. 



160 E. R. PEACOCK 

Celsus Lewis, 1895: 118. (Here transferred from the Dasytinae (Pic, 1937: 56).) Type species: Celsus 

spectabilis Lewis, 1895: 119, by monotypy. Syn. n. 
Eurelymis Casey, 1895: 600. (Here transferred from the Melyrinae (Pic, 1929: 17; Blackwelder, 1975: R67, 

11.) Type species: Eurelymis speculifer Casey, 1895: 603, by PRESENT DESIGNATION. Syn. n. 

Colour varying from black to dark, light, or reddish brown, often with elytra and pronotum different in 
colour or shade, legs sometimes lighter than body. Body clothed with very short, sparse, adpressed 
sometimes scale-like setae. 

Head tending to be longer than broad, retractable under pronotum, frons without lines or depressions 
except for a faint short longitudinal groove mid way between eyes; punctures weakly rimmed, sometimes 
confused. Eyes slightly emarginate when seen from above, glabrous. Antennae short, not usually reaching 
beyond base of pronotum, serrate from segment 4, apical five or six segments often much more broadly 
serrate than the basal segments, giving an almost clubbed appearance (Fig. 1); sexually dimorphic. Apical 
segment of maxillary palps feebly securiform, sides subparallel, sensory surface diagonal. Pronotum 
broader than long, but not markedly so, sides serrate, evenly arcuate and visible from above; narrower 
than base of elytra, without anterior or posterior borders; all angles rounded; punctures rimmed, often 
very dense and confused and becoming tuberculate at sides making margin appear serrated, each tubercle 
with a forwardly directed curved seta; sublateral setae directed towards disc. Scutellum subquadrate. 
Elytra elongate, subparallel-sided, about twice as long as broad; strongly punctured; epipleura very narrow 
at level of hind coxae and continuing as a very narrow line which deflects laterad and evanesces altogether 
at about level of 2nd or 3rd visible abdominal sternite. The female is characterised by the presence of an 
oval, raised, shiny, unpunctured area on the apical third of each elytron (Fig. 6), except in one Japanese 
species, S. elongatus (Nakane). Hind wing with anal cell elliptical (Fig. 58). Tarsi simple, elongate; claws 
small with minute teeth near base (Fig. 5). (Legs very slender.) Prosternal process narrow but extending 
between front coxae to mesosternum when body is contracted. Metasternum convex (Fig. 3). Length 
3-0-6-0 mm. 

Distribution. C. & E. Europe, Asia, Japan, NW. America (including Canada). 

Material examined 

7 species, including types of type species of Celsus (BMNH) and Eurelymis (MCZ) and 3 reliably identified 

specimens of the type species of Semijulistus (2 in MNHN). Also types of Pic's N. American species 

(MNHN). 

Remarks. This genus is included in the Rhadalinae by Majer (1983) although he regards the latter as a 
subgroup of the Dasytinae. 

Although the distribution is so wide and the genus has been described three times, examination of the 
wing venation, mouthparts, genitalia etc. shows that the various elements are congeneric. The main 
difference between the Japanese species and the others is the shape of the apex of the last abdominal tergite 
in the female, but this is believed to be of only specific importance. 

Semijulistus is closely related, and similar in appearance, to Microjulistus Reitter which is mainly from 
the Mediterranean region, Africa and western U.S.S.R. These genera have the basic characters of the 
Rhadalinae, but form a group apart from the other genera. They have a more slender, delicate appearance, 
are generally of smaller size with slender tarsi and recumbent pubescence and differ also in the form of the 
elytral epipleura. Most genera of the subfamily are generally larger and of stouter appearance and have 
longer, semi-erect or suberect setae. 

Semijulistus differs from Microjulistus in its larger size, its unpunctured areas on the elytra of the female 
(in most species), its more strongly punctured pronotum, its lack of a linear lateral edge to the pronotum, 
its more convex metasternum, its apically divided tegmen, the fact that the pronotal sublateral setae point 
towards the disc and its distinct wing venation with an oval anal cell. 

Checklist of species 

ater (LeConte), 1878: 461 (from Eurelymis). N. America, Canada. Comb. n. 
bicoloripes Pic, 1928^(431): 54 hors texte. N. America. 
callosus (Solsky), 1867: 32 (Dosytes). U.S.S.R. (Turkestan). 
elongatus (Nakane) , 1981: 129 (from Celsus). Japan. Comb. n. 
flavipes (LeConte), 1878: 461 (from Eurelymis). N. America. Comb. n. 
rubrithorax Pic, 1928a(431): 54 hors texte. N. America. 
spectabilis (Lewis) , 1895: 119 (from Celsus). Japan. Comb. n. 
speculifer (Casey) , 1895: 603 (from Eurelymis). N. America. Comb. n. 



REVIEW OF THE RHADALINAE 161 

MICROJULISTUS Reitter 

(Figs 2, 4, 59) 

Microjulistus Reitter, 1889a: 111. Type species: Micro julistus fulv us Reitter, 1889a: 112, by monotypy. 
Ceralliscus Bourgeois, 1894: 121. Type species: Ceralliscus raffrayi Bourgeois, 1894: 122, by monotypy. 

Similar to Semijulistus , but smaller. Form delicate, with slender legs and recumbent, often scale-like, 
usually light brown or greyish setae which obscure cuticle. 

Head as in Semijulistus but clypeus rather longer and more parallel-sided, frons without depressions or 
grooves; cuticle rugose, often obscured by setae, punctures obscure. Eyes oval, glabrous. Antennae and 
palps similar to those of Semijulistus, sensory surface of latter more strongly diagonal. Pronotum only 
weakly transverse, about as broad as elytra at base; sides smooth, not or scarcely crenulate, lateral margin 
not visible from above due to strong lateral convexity, but visible from beneath as a fine line, not bordered; 
anterior angles rounded or obsolete, posterior either rounded or obtuse; punctures confused, sometimes 
rimmed, cuticle rugose, mat; sublateral setae directed towards base (Fig. 2). Scutellum subquadrate but 
rounded apically. Elytra elongate, subparallel; sides smoothly curved, not visible from above; punctura- 
tion weak, particularly near apex, usually microreticulate between punctures; epipleura very short and 
narrow, visible from side, deflecting laterad anterior to level of hind coxae. Elytra of female as in male. 
Hind wing with weak venation, no anal cell and very long setae along costal margin (Fig. 59). Prosternal 
process as in Semijulistus. Metasternum flattened, with distinct median longitudinal groove (Fig. 4). Length 
under 3 mm. 

Distribution. Mediterranean, Africa (widespread but localised), Arabia, C. Europe, C. Asia. 

Material examined 

8 named species, including reliably identified specimen of type species, and some material not determined 

at species level. 

Remarks. In Pic's catalogue (1937) this genus was regarded as a dasytine. Majer (1983) included it in the 
Rhadalinae although he regards the latter as a subgroup of the Dasytinae. 

In the examples of Microjulistus examined, the tegmen is entire apically, not divided as it is in 
Semijulistus, and the dorsal appendage is less moveable. 



Checklist of species 

chobauti (Pic), 1895: 79. Algeria. {Ceralliscus) 
duboisi Peyerimhoff , 1931a: 51. Algeria. 
fulvus Reitter, 1889a: 112. U.S.S.R., Mongolia. 

obscurithorax Pic, 1905: 97. Turkestan. 

moreli (Pic), 1927: 10. (Ceralliscus) 

pectoralis (Pic), 1927: 10. (Ceralliscus) 

rufithorax (Pic), 1927: 10. (Ceralliscus) 
gibbipennis Chobaut, 1898: 80. S. Algeria. 
laticollis (Pic), 1894: 95. Algeria. (Ceralliscus) 
lysholmiYic, 1898: 170, 171. Syria. 

abdominalis Pic, 1898: 170, 171. 

nigripennis Pic, 1898: 170, 171. 
me ynieri Peyerimhoff , 1931a: 51. Algeria. 
minutusFic, 1927: 5. Corsica. 
nigricollis (Pic) , 1894: 95. Algeria. (Ceralliscus) 
olivaceusPic, 1931c: 446. Ethiopia. 
raffrayi (Bourgeois), 1894: 122. Algeria. (Ceralliscus) 

nigrifrons (Pic), 1894: 95. (Ceralliscus) 
rubricollis (Abeille) , 1907: XXII [reference not seen]. Arabia. (Ceralliscus) 
subconvexus Pic, 1903a: 179. S. Africa. 

nigricolor Pic, 1903a: 179. 

fulvithorax Pic, 19046: 33. 
wegeneri Pic, 1898: 170. Egypt. 

obscurus Pic, 1898: 170, 171. 



162 E. R. PEACOCK 

Acknowledgements 

I thank the following for the loan of specimens: Dr G. E. Ball, University of Alberta, Edmonton, Canada; 
Dr M. Brancucci, Naturhistorisches Museum, Basel, Switzerland; Dr H. Dybas, Field Museum of Natural 
History, Chicago, Illinois, U.S.A.; Dr L. H. Herman Jr, American Museum of Natural History, New 
York, U.S.A.; DrD. H. Kavanaugh, California Academy of Sciences, San Francisco, California, U.S.A.; 
Dr J. M. Kingsolver, National Museum of Natural History, Smithsonian Institution, Washington D.C., 
U.S.A. ; Dr K. Majer, Faculty of Forestry, University of Agriculture, Zemedelska, Brno, Czechoslovakia; 
Dr J. J. Menier, Museum national d'Histoire naturelle, Paris, France; Dr T. Nakane, Kagoshima 
University, Korimoto, Kagoshima, Japan; Dr Scott R. Shaw and Ms M. K. Thayer, Museum of 
Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S. A. I thank my colleagues Mr 
P. M. Hammond, Dr N. E. Stork, Dr R. B. Madge and Mr R. T. Thompson for reading the manuscript and 
offering helpful criticism and advice and the staff of the Scanning Electron Microscope Unit at the BMNH 
for their help in the production of the photographs. 

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1896. In Kiister & Kraatz, Die Kdfer Europa's 32: nos 1-100, 32A-Q. Niirnberg. 

1897. Trichoceble heydeni n. sp. aus China. Deutsche Entomologische Zeitschrift 1: 155. 
1897a. In Kiister & Kraatz, Die Kdfer Europa's 33: nos 81-100. Niirnberg. 

18976. In Kiister & Kraatz, Die Kdfer Europa's 34: nos 1-100, 34A-34BBB. Niirnberg. 

1900. In Kiister & Kraatz, Die Kdfer Europa's 37: no. 4. Niirnberg. 

1903. In Kiister & Kraatz, Die Kdfer Europa's 40: nos 97, 98. Niirnberg. 

1906. In Kiister & Kraatz, Die Kdfer Europa's 42: nos 99, 100; 43: nos 13-17. Niirnberg. 



Schonherr, C. J. 1817. Synonymia Insectorum 1 (3): iii-xi, 1-266. Skara. 

Solsky, S. 1867. Coleopteres nouveaux. Horae Societatis Entomologicae Rossicae 5: 29-37. 

Stephens, J. F. 1829. A Systematic Catalogue of British Insects. 388 pp. London. 

1830. Illustrations of British Entomology . Mandibulata 3: 1-374, index, pis 16-19. London. 

Suffrian, E. 1843. Entomologische Bemerkungen. Stettiner Entomologische Zeitung 4: 330-337. 

Thomson, C. G. 1859. Skandinaviens Coleoptera 1: 290 pp. Lund. 

Vinson, J. 1946. The Dasytidae of the Mascarene Islands. Mauritius Institute Bulletin 2: 249-277. 

1953. Coleopteres nouveaux ou peu connus de l'ile de la Reunion. Bulletin de la Societe Entomolo- 

gique de France 58: 143-146. 
1957. On two species of Pelecophora from Round Island with the description of a larva of this genus 

(Col. Dasytidae). Mauritius Institute Bulletin 5: 1-6. 
1958. Catalogue of the Coleoptera of Mauritius and Rodriguez, Part 2. Mauritius Institute Bulletin 4: 



75-130. 

1967. Liste chorologique des Coleopteres des Mascareignes. Mauritius Institute Bulletin 4: 299-394. 



Waterhouse, C. O. 1876. New species of Coleoptera from the Island of Rodriguez, collected by the 

naturalists accompanying the Transit-of- Venus Expedition. Annals and Magazine of Natural History (4) 

18:105-121. 
Watt, J. C. 1975. Notes on priority of family-group names in Coleoptera. The Coleopterists Bulletin 29: 

31-34. 
Wittmer, W. 1935. Risultati Scientifici delle cacce entomologiche di S.A.S. il Principe Alessandro della 

Torre e Tasso. II. Malacodermata. Bollettino de R. Laboratorio di Entomologia Agraria di Portici 28: 

247-256. 
1938. Zur Kenntnis der Malacodermata Aegyptens. Bulletin Societe Entomologique d'Egypte 21 

(1937): 167-170. 
1940. Neue Malacodermata aus der Sammlung des Museum Heude, Shanghai. Notes d' Entomologie 

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1941. Malacodermata von den Philippinen aus der Sammlung des zoologischen Museums in 

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summary.] 



Index 



Synonyms are in italics; 
references to descriptions, 
position in key and in 
checklists are in bold. 

abdominalis Pic 161 
abietum Kiesenwetter 153 
acutangulus Schilsky 153 
adaliensis Pic 155 
adanensis Pic 153 
aerosus Schilsky 153 
aestivus Kiesenwetter 153 



aestivus Kiesenwetter 154 
afer Schilsky 153 
ahenus Kiesenwetter 157 
akbesianus Pic 153 
albipilis Kiesenwetter 153 
albonotata Pic 147 
albovillosa Vinson 147 
alfierii Wittmer 156 
algiricus Schilsky 153 
alluaudi Pic (Aplocnemus) 153 
alluaudi (Pic) (Malthacodes) 150 
alpestris Kiesenwetter 153 



alternatus Peyerimhoff 153 
ambrensis (Pic) 150 
anatolicus Schilsky 153 
andalusicus (Rosenhauer) 153 
angolanus Wittmer 153 
angustata Pic 147 
antelmei Alluaud 147 
Anthriboclerus Schenkling 130, 

131, 132, 136, 139, 143, 144, 

145, 146, 147, 148 
Antinea Peyerimhoff 129, 131, 

132, 136, 158, 159 



168 



INDEX 



Aplocnemina 131 
Aplocneminae 129, 130, 131 
Aplocnemus Stephens 129, 131, 

132, 135, 136, 138, 140, 143, 

149, 150, 151, 152, 153, 158 
araratica Yablokov-Khnzoryan 

158 
arbustorum (Kiesenwetter) 158 
areschanus Pic 154 
ater (LeConte) 132, 134,141, 

160 
atra LeConte 132 
atricolor Pic 147 
atricornis Pic 153 
aubei Kiesenwetter 153 

baborensis Pic (1908) 153 
baborensis Pic (1922) 153 
barklyi Vinson 147 
barnevillei Kiesenwetter 153 
basalis (Kuster) 153 
beauprei Pic 153 
bequaerti (Pic) 150 
berytensis Sahlberg 153 
bicoloratus Pic 153 
bicoloripes Pic 160 
bimaculatus (Pic) 150 
biscrensis Schilsky 153 
biskraensis Schilsky 153 
biskrensis Schilsky 153 
bonnairei Pic 156 
borbonica Vinson 147 
bourbonicus (Pic) 150 
bourgeoisii (Fairmaire) 150 
brevis (Pic) (Malthacodes) 150 
brevis (Rosenhauer) 
(Aplocnemus) 153 
brevissimus Pic 153 
breviusculus Schilsky 153 
brunnescens Pic 155 
brunneus (Pic) 150 

caelatus Brulle 153 
caerulescens Schilsky 156 
calidus Mulsant & Rey 153 
callosus (Solsky) 159, 160 
capillicornis Abeille 153 
capraianus Pic 155 
caramanicus Sahlberg 153 
cariei Pic 147 

caroli Pic (Aplocnemus) 153 
caroli (Pic) (Malthacodes) 150 
castaneus (Gorham) 145 
castiliensis Schilsky 153 
caucasicus Schilsky 153 
Celsus Lewis 129, 131, 132, 133, 

160 
Ceralliscus Bourgeois 132, 161 
chalconatus (Germar) 154 
chalybeus (Germar) 155 
charmoyi Alluaud 147 
chlorosoma (Lucas) 154 
chobauti (Pic) 161 
cincta Pic (Trichoceble) 158 
cinereovariegatus Blair 150 
cioides (Champion) 149, 150 



Cleridae 130, 146 
Cleroidea 129 
cobosi Pic 154 

coeruleatus (Rosenhauer) 154 
colasi Pic 154 
concinna Vinson 147 
conradsi (Pic) 150 
consobrinus (Rosenhauer) 154 
convexa Pic 158 
corcyricus Miller 154 
Corynetinae (Cleridae) 146 
crenicollis Kiesenwetter 154 
cribrarius (Brulle) 154 
cribricollis Mulsant & Rey 154 
cribripennis Pic 154 
cribripennis Pic 153 
cribrosus Schilsky 154 
croceicornis Kiesenwetter 154 
cuprea (Lucas) 154 
cupreatus Schilsky 154 
curticornis Pic 154 
curtipennis Pic 154 
curta (Baudi) (Trichoceble) 158 
curtus Pic (Aplocnemus) 154 
cyaneus Champion 145 
cylindricus Kiesenwetter 154 
Cymbolus Gorham 129, 131, 132, 

144, 145 
cyrenaicus Pic 154 

darutyi Pic 147 

Dasyrhadus Fall 144 

Dasytes 151, 152, 153, 154, 155, 

156, 157, 159, 160 
Dasytinae 129, 130, 147, 150, 151, 

153, 158, 159, 160, 161 
decorata Pic 147 
delagrangei Pic 154 
dentatus Schilsky 154 
depressicollis Schilsky 154 
desertorum Pic 154 
diaphanus Schilsky 154 
difficilis Holdhaus 154 
Diglobicerus Latreille 132, 146 
Diplambe Schilsky 131, 132, 136, 

152, 153 
disconotatus (Pic) 150 
distinctipes Pic 153 
distinctus (Fairmaire) 150 
diversus Schilsky 155 
Donaldia Alluaud 131, 132, 149, 

150, 151 
duboisi Peyerimhoff 161 
duplicatus Kiesenwetter 154 

edoughensis Pic 156 

elegans (Alluaud) (Malthacodes) 

150 
elegans Majer (Kubanius) 132, 

135, 142, 143, 149, 157 
Elicopis Stephens 132, 151, 152 
elongata Schilsky (Trichoceble) 

158 
elongatior Pic 155 
elongatus (Champion) (Rhadalus) 

132, 143, 145 



elongatus (Nakane) (Semijulistus) 

160 
elongatus (Pic) (Malthacodes) 150 
emmerezi Pic 147 
erosus Mulsant & Rey 155 
escalerai Pic 154 
Eucymbolus Champion 131, 132, 

136, 144, 145 
eumerus Mulsant & Rey 154 
Eurelymis Casey 129, 131, 132, 

133, 160 

fairmairei (Alluaud) 150 

fasciatus (Pic) 150 

fauconneti Pic 154 

femoralis Illiger 155 

femoralis Schilsky 155 

femoratus Schilsky 155 

festai Pic 154 

flavicornis Schilsky 154 

flavipes (LeConte) (Semijulistus) 

132, 143, 160 
flavipes Schilsky (Aplocnemus) 

156 
floralis (Olivier) 135, 138, 141, 

143, 158 
floricola (Kiesenwetter) 158 
florigersp. n. 129, 135, 139, 141, 

142, 143, 147, 148 
fortepunctatus Pic 154 
fulvipes Schilsky 156 
fulvithorax Pic 161 
fulvohirta (Brisout de Barnville) 

159 
fulvopilis (Reitter) 158 
fulvus Reitter 134, 161 
funera (Kiesenwetter) 158 
fuscitibia Mulsant & Rey 155 



gedyei (Pic) 150 
geniculatus Schilsky 154 
gestroi Schilsky 154 
gibbipennis Chobaut 161 
gracilicornis Schilsky 154 
grancanariensis Lindberg 154 
grandis Schilsky 158 
gridellii Pic 153 
griseohirta (Reitter) 158 
griseopubescens Pic 154 

hamoni Vinson 147 
Haplocnemates Mulsant & Rey 

131 
Haplocneminae Crowson 131 
Haplocnemus; Agassiz 132, 152 
hebraicus Schilsky 154 
Helicopis; Agassiz 132, 152 
Helicopis Fabricius (Lepidoptera) 

152 
Hellicopis Fabricius 152 
Hemipleurusgen. n. 129, 131, 

132, 135, 136, 139, 141, 142, 

143, 144, 147, 148, 157 
heydeni Schilsky (Aplocnemus) 

154 



INDEX 



169 



heydeni Schilsky (Trichoceble) 

158 
hickeri Pic 154 
hierichunticus Sahlberg 154 
hispanicus Pic 154 
Holcopleura Schilsky 132, 152 
holtzi Pic 153 

illigeri (Gyllenhal) 135, 138, 140, 

142, 143, 146, 147 
immaculatus (Pic) 150 
immarginata Reitter 159 
imperforatus Pic 154 
impressa Marsham 151, 152, 154 
impressipennis Pic 154 
impressus (Marsham) 135, 138, 

154 
incognitus (Faldermann) 154 
indicus (Champion) 129, 149, 150, 

153 
Indiodasytes Pic 129, 130, 131, 
v 132, 137, 138, 141,142, 143, 151 
interrupta Alluaud (Pelecophora) 

147 
interrupta (Pic) (Malthacodes) 151 
integer Baudi 154 
Ischnopalpus Schilsky 131, 132, 

140, 152, 153 

jeanneli Pic 147 
jejunus Kiesenwetter 154 
Julistus Kiesenwetter 131, 133, 
158, 159 

kaszabi Majer 155 
kiesenwetteri Schilsky 155 
korbi Schilsky 155 
koziorowiczi Desbrochers des 

Loges 155 
krugeri Pic 155 
kubanensis Pic 155 
Kubanius Majer 131, 132, 135, 

136, 142, 148, 157, 158 
kuesteri Schilsky 156 

lacoi Majer 155 
laeta Majer 159 
laetipes Abeille 155 
laetus Schilsky 153 
Lagria Fabricius 152, 155 
lateralis Schilsky 155 
laterufus (Pic) 150 
latesuturalis (Pic) 150 
laticollis (Pic) 161 
latior Pic 155 
latipennis Pic 155 
lecontei Casey 145 
lederi Schilsky 159 
libanicus Pic 155 
limbipennis Kiesenwetter 155 
lixensis Bourgeois 156 
longicornis (Kiesenwetter) 

(Trichoceble) 159 
longicornis Kocher (Aplocnemus) 

155 
longior Fall 144 



longulus Schilsky 155 
luteofasciatus (Pic) 150 
lysholmi Pic 161 

macedonicus Pic 155 
maculatipennis (Pic) 150 
maculatus (Pic) (1906) 150 
maculatus (Pic) (1926) 150 
madurensis (Pic) 137, 138, 142, 

151 
maindroni (Pic) 149, 150 
major Pic 159 
Malachiinae 129 
Malthacodes Waterhouse 129, 

131,132,135,140,141,142, 

143,147,148,149,150,151,153 
mancinii Pic 155 
marchali Pic 154 
marginalis Fairmaire 147 
marginatus Rottenberg 156 
marginicollis Schilsky 155 
martini (Fairmaire) 150 
mateui Pic 156 
maurus Schilsky 155 
mediocris Majer 159 
melitensis Schilsky 155 
Melyridae Leach 129, 130, 131, 

144 
Melyrinae 129, 130, 144, 145, 160 
Melyris Fabricius 130, 158 
memnonia (Kiesenwetter) 158, 

159 
metallicus (Pic) 150 
meynieri Peyerimhoff 161 
Microjulistus Reitter 130, 131, 

132, 133, 134, 143, 152, 160, 161 
minor (Pic) 150 
minutus (Pic) (1906) 

(Malthacodes) 150 
minutus (Pic) (1931) 

(Malthacodes) 129, 150 
minutus Pic (Microjulistus) 161 
mirabilis Schilsky 155 
moestus (Gorham) 129, 149, 150, 

153 
mohamed Chobaut 155 
montivagus (Rosenhauer) 152, 

155 
moreli (Pic) 161 
morio (Schonherr) 155 
multisignata Pic 147 

nevadensis Pic 155 

nigra Pic 159 

nigrescens Pic 153 

nigriceps (Pic) 150 

nigricollis (Pic) 161 

nigricolor (Pic) (Malthacodes) 150 

nigricolor Pic (Microjulistus) 161 

nigricornis (Fabricius) 140, 143, 

152, 155 
nigrifrons (Pic) 161 
nigripennis Pic 161 
nigripes Mulsant & Rey 155 
nigrolineata Guerin 147 
nigromaculatus (Pic) 150 



notaticeps (Pic) 150 

obliquata Alluaud 147 

obscuricollis Pic 147 
obscuripes Schilsky 156 
obscurithorax Pic 161 
obscurus Germar (Aplocnemus) 

156 
obscurus (Pic) (Malthacodes) 150 
obscurus Pic (Microjulistus) 161 
ocularis Reitter 159 
oculata Schilsky 159 
oertzeni Schilsky 158, 159 
olivaceus Pic 161 
orientalis Lohse 157 
oxylepisiformis (Pic) 150 

palaestinus Baudi 153, 155 
pallidipes (Pic) 159 
pallidithorax (Pic) 150 
pandellei Pic 156 
parumpunctatus Schilsky 155 
parvus nom. n. 129, 150 
pectinatus (Kiister) 155 
pectinicornis (Dufour) 153 
pectinicornis (Lucas) 155 
pectoralis (Pic) 161 
Pelecophora Dejean 131, 132, 

135, 138, 140, 142, 143, 146, 147 
Pelecophorus Berthold 146 
pellucens Kiesenwetter 155 
perforatus (Pic) (Malthacodes) 

150 
perforatus Schilsky (Aplocnemus) 

155 
perrieri (Fairmaire) 150 

persicus Schilsky 155 

pertusus Kiesenwetter 155 

pesruchesi Schilsky 155 

peyroni Pic 155 

philippinus Wittmer 155 

Phycosecidae 130 

pictus Waterhouse 135, 142, 143, 

149, 150 
pikei Vinson 131, 147 
pini (Redtenbacher) 154 
pinicola Kiesenwetter 154, 156 
ponferradanus Pic 155 
Prionocerinae 129 
pristocerus Kiesenwetter 155 
Pseudaphyctus Pic 132, 152 
pubicornis Wittmer 151 
pulverulentus (Kiister) 156 
punctatus (Germar) 155 
puncticollis Sahlberg 156 
punctiger Schilsky 156 
punctipennis (Gorham) 145 

quadrituberculatus (Champion) 

145 
quercicola Mulsant & Rey 156 

raffrayi (Bourgeois) 

(Microjulistus) 161 
raffrayi Pic (Aplocnemus) 156 



170 

ragusae Schilsky 156 
ramicornis Kiesenwetter 

(Aplocnemus) 156 
ramicornis Schilsky (Trichoceble) 

159 
raymondi Deville 156 
rechmayanus Pic 155 
reitteri Schilsky 152, 156 
Rhadalidae 130, 131 
Rhadalini LeConte 131, 144 
Rhadalinae LeConte 129, 130, 

131, 132, 133, 144-148, 150, 

151,153,157-161 
Rhadalus LeConte 129, 130, 131, 

132, 136, 137, 138, 141, 143, 

144, 145, 152 
robustior Pic 154 
rubricollis (Abeille) 161 
rubrithorax Pic 160 
rubronotatus (Pic) 151 
ruficollis (Pic) 151 
rufipes Miller 156 
rufitarsis (Sahlberg) 156 
rufithorax (Pic) (Malthacodes) 150 
rufithorax (Pic) (Microjulistus) 

161 
rufofemoratus Pic 154 
rufolateralis Pic 155 
rufomarginatus Perris 156 
rufomarginatus Schilsky 155 
rufopiceus (Gorham) 137, 138, 

141,144,145 
rugicollis Schilsky 156 
rugulosus (Rosenhauer) 156 
russicus Reitter 156 

sanctus Pic 140, 156 
saxicola Peyerimhoff 159 
schatzmayeri Pic 159 
schilskyi Reitter 159 
scotti Schenkling 139, 143, 145, 

146 
sculpturatus Wollaston 156 
semicaeruleus Pic 156 
Semijulistus Schilsky 129, 131, 



INDEX 

132,133,134,141,143,152, 

159, 160, 161 
semirufescens (Pic) 159 
serbicus Kiesenwetter 156 
serratus (Brulle) 156 
serratus Redtenbacher 154 
serrulatus Schilsky 156 
sicardi (Pic) 151 
siculus Kiesenwetter 156 
similis Schilsky 156 
sinuatus (Pic) 151 
smyrnensis Pic 156 
sparsepunctata Pic 159 
spectabilis (Lewis) 134, 160 
speculifer (Casey) 160 
strandi Marcu 156 
subacuminatus Pic 156 
subapicalis (Pic) 151 
subcaerulea Pic (Trichoceble) 159 
subcoeruleus Pic (Aplocnemus) 

156 
subconvexus Pic 143, 161 
subcoriacea Reitter 159 
subcostatus Schilsky 152, 153, 156 
subdepressus (Pic) 151 
subelongatus Pic 153 
subfasciatus (Pic) 151 
subglabra Alluaud 147 
subinteger Pic 156 
subviolaceus Pic 153 
suggara Peyerimhoff 156 
suturalis (Pic) 151 
syriacus Schilsky 156 

tarsalis (Sahlberg) 156 
tarsalis Schilsky 153 
tarsicola Sahlberg 156 
taygetanus Pic 156 
tenerifensis Pic 156 
testaceipes Pic (Aplocnemus) 156 
testaceipes Pic (Trichoceble) 159 
testaceofemoralis Pic 155 
testaceus LeConte 138, 144, 145 
thalensis Pic 156 
theresae Pic 153 



theryi Pic 153 
thessalicus Pic 156 
tibialis Schilsky 157 
tibiellus Schilsky 156 
tingitanus Baudi 156 
torretassoi Wittmer 159 
tournieri Pic 152, 157 
Trichoceble Thomson 131, 132, 

133,135,136,138,141,143, 

152, 157, 158, 159 
trimaculata Pic 147 
trinacriensis Ragusa 157 
trinotatus (Pic) 151 
tuberculifer Motschulsky 157 
tumidus Kiesenwetter 152, 157 
tuniseus Pic 157 
turcicus Schilsky 157 

uhagoni Schilsky 157 
unguicularis Reitter 159 
unicolor Pic 159 
unimaculatus (Pic) 150 

vageguttatus (Fairmaire) 140, 142, 

143, 149, 150, 151 
variabilis (Pic) 151 
variolatus Costa 155 
venustulus Schilsky 157 
vestitus Wollaston 157 
viberti Pic 156 
vicinus (Pic) 151 
viertli Schilsky 155 
vinsoni Pic 147 
virens (Kuster) 156 
virens (Suffrian) 157 
viridescens Pic 157 
vittata Laporte de Castelnau 147 

wegeneri Pic 161 
wolcotti (Hinton) 132, 145 

Xamerpus Fairmaire 129, 131, 

132, 149, 150, 151 
xanthopus Kiesenwetter 155 
xanthopus Schilsky 156 



British Museum (Natural History) 

Milkweed butterflies: their cladistics and biology 

P. R. Ackery & R. I. Vane- Wright 

The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of 
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courtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry and 
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Started in 1976 within the conventions of evolution by natural selection and Hennig's 
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425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. 
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Titles to be published in Volume 56 



The legume-feeding psyllids (Homoptera) of the west Palaearctic Region 

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A review of the Malvales-feeding psy Hid family Carsidaridae (Homoptera) 

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Bulletin of the 

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A revision of some Afrotropical genera of 
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J. Quinlan 






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Entomology series 

Vol 56 No 4 28 April 1988 



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Entomology series 
Vol 56 No 4 pp 171-229 



Issued 28 April 1988 



A revision of some Afrotropical genera of Eucoilid ae 
(Hymenoptera) 

!rn ' 

J. Quinlan. ^ 

Department of Entomology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Contents 

Synopsis 171 

Introduction 171 

Material examined and terminology 172 

Depositories 172 

Classification 172 

Checklist of Afrotropical Eucoilidae included in this paper 173 

Amendments to key to genera of Afrotropical Eucoilidae 173 

Angustacorpa gen. n 174 

Bothrochacis Cameron 176 

Diglyphosema Foerster 179 

EmargoWeld 181 

Eucoila Westwood 186 

Glauraspidia Thomson 189 

Leptopilina Foerster 191 

Sirenes gen. n 201 

Acknowledgements 205 

References 206 

Index 207 



Synopsis 

Eight genera comprising 50 Afrotropical species are revised. Two genera {Angustacorpa and Sirenes) and 
40 species are newly described; the primary types of 14 nominal species have been examined. Amendments 
to the key to genera (Quinlan, 1986) and a checklist are included. 

Introduction 

In this second part of a revision of the Afrotropical Eucoilidae a further 8 genera and 50 species 
are dealt with, including 2 new genera and 40 new species. My key to genera (Quinlan, 1986) has 
been amended to include the newly described Angustacorpa and Sirenes, and Emargo Weld, 
here transferred from the Figitinae. During* the preparation of this paper all accessible type 
specimens have been examined and redescribed. When type material was not available the 
position of the nominal species concerned is discussed in relation to other taxa. 

Weld (1960) originally placed Emargo in the subfamily Figitinae and stated that it differed 
from all other figitines in having the fore wing deeply emarginate. This character was otherwise 
known to occur only in the eucoilid genus Kleidotoma. Examination of the type species Emargo 
eciton shows it to possess a scutellar cup, although this is not of the usual form (Fig. 51). Weld's 
placement of the genus was influenced by the relative lengths of the gastral tergites (i.e. tergite 2 
shorter than 3, see discussion p. 181). 

Apart from the eight genera dealt with here and the ten dealt with in my earlier paper 
(Quinlan, 1986) there are at least three further genera of Eucoilidae (Trybliographa, Daruna 
and Ganaspis) in the Afrotropical region. 



\A'OS. 



Bull. Br. Mus. Nat. Hist (Ent.) 56 (4) 171-229 Issued 28 April 1 ! 



172 J. QUINLAN 

Material examined and terminology 

The material on which this paper is based has come from the same sources as that included in my 
two previous papers on the Afrotropical Cynipoidea (Quinlan, 1979, 1986). Terminology is as 
indicated in those papers. 

Depositories 

Type depositories are abbreviated in the text as follows. 

AM Albany Museum, Grahamstown 

BMNH British Museum (Natural History), London 

MA Musee d' Amiens, Amiens 

MNHN Museum National d'Histoire Naturelle, Paris 

MNHU Museum fur Naturkunde der Humboldt Universitat, Berlin 

MRAC Musee Royale de TAfrique Centrale, Tervuren 

SAM South African Museum, Cape Town 

USNM United States National Museum of Natural History , Washington , D . C . 

ZI Zoologiska Institution, Lund 

ZSBS Zoologische Sammlung des Bayerischen Staates, Munich 

Classification 

Reference to my earlier paper (Quinlan, 1986) will aid appreciation of taxonomy, classification 
and relationships of the genera dealt with here. The affinities of Bothrochacis and Eucoila are 
also treated by Nordlander (1981, 1982). Diglyphosema is placed by Nordlander (1982) in the 
Gronotoma-group of genera (Nordlanderia etc). Excepting Zaeucoila, a Neotropical genus not 
examined by me, I agree that Diglyphosema is related to this group. All five genera have a 
plesiomorphic form of pronotal plate (i.e. it does not project forward) which is similar to that 
found in the Figitinae and the cynipine genera Aulacidea and Ay lax. I have examined 
Amphiglyphosema, a genus erected by Benoit in 1956, and consider it to be congeneric with 
Diglyphosema, thus making it the first record of a Diglyphosema from the Afrotropical region. 
Glauraspidia is placed by Nordlander (1982) in the Chrestosema-group, comprising Chres- 
tosema, Glauraspidia, Pseudopsichacra, Odonteucoila, Dieucoila, and Leptolamina. At present 
two species of Glauraspidia, newly described here, are known from the Afrotropical region. The 
new genus Sirenes is very closely related to Glauraspidia but differs in the sculpture of the head, 
mesoscutum and mesopleuron, the radial cell of the fore wing and the form of the antenna in 
both sexes. The characters common to both are the absence of a complete mesopleural suture, 
the dense pubescence on either side of the pronotal plate and entire metapleuron, the density of 
the pubescence on the propodeal area and the dense ring of felt-like pubescence at the base of 
segment one of the gaster. 

Angustacorpa, newly erected, has a projecting pronotal plate with the lateral foveae between 
the anterior and posterior parts fused or closed (Fig. 11) as in Stentorceps, Rhoptromeris and 
Trichoplasta. It is distinguished, however, on the strongly compressed head and thorax (Fig. 4) 
and the position of the antennal sockets (Fig. 1). The genus Leptopilina has been the subject of a 
cladistic analysis by Nordlander (1980). He considers it to belong to the Rhoptromeris-group of 
genera comprising Rhoptromeris, Cothonaspis, Trichoplasta and Leptopilina. Leptopilina and 
Cothonaspis have very similar pronotal plates (i.e. the foveae on either side of the medial bridge 
are open on the lateral margins and not fused laterally as in Trichoplasta and Rhoptromeris). 
Leptopilina has a short thorax, the compound eyes large and the head constricted behind the 
eyes. Cothonaspis has an elongate slender thorax and the compound eyes are small. Emargo, as 
stated above, is considered for the first time to be a eucoilid and is transferred from the Figitinae 
(seep. 181). 



AFROTROPICAL EUCOILIDAE 



173 



Checklist of Afrotropical Eucoilidae included in this paper 



EUCOILIDAE Thomson, 1862 
ANGUSTACORPA gen. n. 
apsus sp. n. 
persa sp. n. 
prodicus sp. n. 
triton sp. n. 
BOTHROCHACIS Cameron, 1904 

Stirenocoela Cameron, 1910 

Ditrupaspis Kieffer, 1910 

Salpictes Kieffer, 1913 
erythropoda Cameron, 1904 

striaticollis (Cameron, 1910) 

semirufa (Kieffer, 1910) syn. n. 

stercoraria Bridwell, 1919 syn. n. 
rufiventris (Kieffer, 1913) 
serratepilosa Benoit, 1956 
DIGLYPHOSEMA Foerster, 1869 

Amphiglyphosema Benoit, 1956 syn. n. 
latesulcatum (Benoit, 1956) comb. n. 
utica sp. n. 
EMARGO Weld, 1960 
ascia sp. n. 
cantos sp. n. 
capito sp. n. 
laverna sp. n. 
matius sp. n. 
micipsa sp. n. 
numa sp. n. 
pallor is sp. n. 
peleus sp. n. 
pexus sp. n. 
the mis sp. n. 
vacuna sp. n. 
EUCOILA Westwood, 1833 

Psilodora Foerster, 1869 

Lytosema Kieffer, 1910 

Psilodoropsis Hedicke, 1913 
bantia sp. n. 



camerounensis Risbec, 1956 
erinna sp. n. 
marina sp. n. 
ve/edasp. n. 
GLAURASPIDIA Thomson, 1862 

Aglaotoma Foerster, 1862 

Apistophyza Foerster, 1869 

Diranchis Foerster, 1869 
casca sp. n. 
scyphus sp. n. 
LEPTOPILINA Foerster, 1869 
apella sp. n. 

atraticeps (Kieffer, 1911) 
oou/ardi (Barbotin, Carton & 

Kelner-Pillault, 1979) 

mahensis (Kieffer, 1911) (Charips) 
secondary homonym 
fannius sp. n. 
faunussp. n. 

heterotoma (Thomson, 1862) 
jfyssp. n. 

mahensis (Kieffer, 1911) (Erisphagia) 
mi sens us sp. n. 
pisonis sp. n. 
syphax sp. n. 
thetus sp. n. 
veste sp. n. 

victoriae Nordlander, 1980 
S/flENESgen.n. 
floccus sp. n. 
orbilussp. n. 
silenus sp. n. 
sinis sp. n. 
sp/o sp. n. 
steropes sp. n. 
sj/mxsp. n. 
sjrtes sp. n. 



Amendments to key to genera of Afrotropical Eucoilidae 

The three genera here added to the Afrotropical fauna can be accommodated in the key to genera 
(Quinlan, 1986) as follows. 

At couplet 8: 
8 Pronotal plate viewed frontally with narrow or obsolete medial bridge, foveae closed 
laterally (Fig. 11). 

Scutellar disc sharply conical, sometimes almost spine- or beak-shaped, overhanging 
propodeum (Fig. 10); $ antenna clavate-subclavate or with segment 4 longer than 3, 
weakly curved on inner margin, sometimes swollen distally; c? antenna with segment 3 
shorter than 4, 4 longer than each of the following segments; radial cell of fore wing open or 

closed (Figs 13, 14) 8A 

Pronotal plate viewed frontally with a wide medial bridge, lateral foveae open, not fused 

externally (e.g. Fig. 27) 9 

8 A Head and thorax viewed dorsally strongly compressed laterally; head in lateral view 
wedge-shaped, in front view with antennae set at the middle (Fig. 1). 

ANGUSTACORPA gen. n. 



174 J. QUINLAN 

Head and thorax viewed dorsally not laterally compressed; head not wedge-shaped in lateral 
view, antennae set above the middle (between the upper margins of the eyes) (Fig. 2). 

TRICHOPLASTA Benoit 

At couplet 16: 

16 Fully winged or brachypterous, apex of wings either incised, arcuate or truncate, not 

rounded, cubitus (M) not distinct, radial cell open on wing margin; tergite 2 of gaster either 
shorter or longer than tergite 3 in lateral view (Figs 22, 52) 16A 

- Fully winged, never brachypterous, apex of wing rounded, rarely truncate, never incised, 

radial cell of fore wing open or closed; tergite 2 of gaster longer than 3 in lateral view (e.g. 

Fig. 193) 17 

16A Tergite 2 of gaster shorter than tergite 3 (Fig. 52); fore wings strongly incised, radial cell open 
on wing margin, veins thick (cf. Figs 65, 67); scutellar cup weakly defined (cf. Fig. 60); 
mesopleural furrow absent EMARGO Weld 

- Tergite 2 of gaster the largest in lateral view (Fig. 22); fore wings either incised, arcuate or 

truncate, radial cell distinctive, vein Ri only thickened at apex, near margin of wing, Rs+M 
usually absent (Fig. 20) or wings short not extending to apex of gaster; scutellar cup distinct 
but small, not extending to apex of longitudinally striate disc which is either rounded or 
conical, spine- or beak-shaped; mesopleural suture percurrent. 

KLEIDOTOMA Westwood 

17 Mesopleural suture either absent or indistinct (Fig. 182); frons prominent in cf , less so in $ ; 

wings in cf of normal length (generally shortened in British specimens); scutellar cup 
raised above level of scutellar disc which is striate to reticulate-rugose; sides of pronotal 
plate and propodeum with dense tufts of pubescence (usually white); tergite 2 of gaster 
with dense felt-like ring of pubescence ; antenna of $ filiform or subclavate 17 A 

- Without the above combination of characters; mesopleural suture generally present; pro- 

notum and propodeum without dense felt-like tufts; tergite 2 of gaster with hairy ring 
variable in density of pubescence; $ antenna usually clavate, cf filiform, 3rd and 4th 

segments sometimes modified 18 

17 A Head, mesoscutum and mesopleuron finely sculptured; radial cell of fore wing completely 
closed on margin (Fig. 188); pronotal plate viewed dorsally, rectangular (Fig. 191). 

SIRENESgen.n. 

- Head, mesoscutum and mesopleuron smooth and polished; radial cell of fore wing generally 

open on margin (Fig. 96); pronotal plate rounded on margins (Fig. 95). 

GLA URASPIDIA Thomson 



ANGUSTACORPA gen. n. 

Type species: Angustacorpa apsus sp. n. 

Diagnosis. $ antenna 13-segmented, clavate and set medially between the eyes; cf antenna 15- 
segmented, filiform with a modified third segment (Fig. 3). Head viewed laterally wedge-shaped, viewed 
dorsally strongly compressed; face long and narrow, eyes closer together than the height of an eye 
measured medially, face smooth and polished; malar groove distinct; ocellar region with long scattered 
hairs (Fig. 1). Pronotal plate with enclosed lateral foveae, one on either side of the medial bridge. 
Mesoscutum smooth and polished without trace of notauli, strongly compressed laterally; scutellar foveae 
polished, round, lateral bars smooth; scutellar disc long, conical, apically rounded, surface reticulate- 
rugose, overhanging propodeum; scutellar cup elliptical, not extending to apex of disc, apical margin of 
cup with a fovea, outer rim of cup with a number of small pits or foveae (Fig. 4). Mesopleuron smooth and 
polished, mesopleural suture complete; metapleuron weakly ridged, anteroventral cavity pubescent; 
propodeum coarsely sculptured on dorsal and lateral surfaces. Segment 1 of gaster short and crenulate, 
obscured by a ring of dense pubescence at base of tergite 2; segments 2-5 of gaster visible in lateral view, 
tergite 2 the largest (Fig. 5). Legs slender, fore and mid coxae hairy. Wings long and narrow, surfaces 
pubescent, margins with a fringe of hairs, radial cell of fore wing long and narrow, completely closed on 
wing margin (Fig. 14). 

Distribution. Kenya, Zaire. 



AFROTROPICAL EUCOILIDAE 175 

Key to species of Angustacorpa gen. n. 

Females 

1 Antennal club 6-segmented (Fig. 6) (distinct, rhinaria pronounced). 

Apex oftergite 2 sparsely punctate, tergites 3-5 finely punctate (Fig. 5) apsussp. n.(p. 175) 

- Antennal club 7-8 segmented (Figs 7, 8) 2 

2 Antennal club 7-segmented (distinct, rhinaria weakly indicated); radial cell of fore wing open 

(Fig. 13). 
Apex oftergite 2 and tergites 3-5 strongly and densely punctate (Fig. 12). ... tritonsp. n.(p. 176) 

- Antennal club 8-segmented; radial cell of fore wing closed (cf . Fig. 14) 3 

3 Antennal segment 3 longer than 4+5, 4 twice length of 5, club segments 3 times as long as wide, 

club very sharply delimited (Fig. 8), apical segments pale yellow prodicus sp. n. (p. 175) 

- Antennal segment 3 shorter than 4+5, 4 longer than 5, club weakly delimited (Fig. 9), apical 

segments of antenna dark, basal segments yellow persa sp. n. (p. 175) 

Angustacorpa apsus sp. n. 

(Figs 4-6, 14) 

Description. $ antenna with segment 3 subequal to 4+5, segments 8-13 with rhinaria, forming a club 
(Fig. 6). Head with long scattered setae; malar grooves percurrent; mandibles tridentate, produced 
forward in lateral view (cf. Fig. 1); occipital carina distinct on lower margins; ocelli widely spaced in dorsal 
view. Pronotal plate viewed dorsally with a broad medial bridge (cf. Fig. 11). Scutellar disc tapering 
apically, not spine-shaped; rim of scutellar cup with six foveae (Fig. 4). Gaster polished, apex of segments 
2-5 punctate (Fig. 5). Wing (Fig. 14). Colour: antenna yellow, club segments slightly darker; head and 
thorax dark chestnut brown; gaster dark dorsally, orange-yellow laterally; legs orange-yellow. 
Cf unknown. 

Material examined 
Holotype $ , Kenya: 15 mis NE. Kisumu, nr Lake Victoria, xi.1979 (Croft) (BMNH). 
Paratypes. Zaire: 2 $ (MRAC). 

Remarks. Separated from the three other species by the sharply delineated 6-segmented antennal club 
(Fig. 6). 

Angustacorpa persa sp. n. 

(Figs 9-11) 

Description. $ antenna with segment 3 shorter than 4+5,4 longer than 5, segments 6-13 with rhinaria, 
forming a weak club (Fig. 9). Head with long lateral setae; malar grooves percurrent; mandibles tridentate, 
produced forward in lateral view (cf. Fig. 1); occipital carina distinct on lower margins; ocelli widely spaced 
in dorsal view. Pronotal plate viewed fronto-dorsally with a broad medial bridge (Fig. 11). Scutellar fovea 
kidney-shaped; scutellar disc beak-shaped, extending past propodeum; scutellar cup long and broad 
medially, rim of scutellar cup with a few small pits or foveae (Fig. 10). Metapleuron strongly ridged. Gaster 
segment 1 as broad as long; tergite 2 with a sparse basal hairy ring; tergites 2-4 visible in lateral view; apex 
of segment 2 and visible parts of 3 and 4 punctate (cf. Fig. 5). Wing (cf. Fig. 14). Coxa and femur with 
scattered long hairs. Colour: antenna yellow basally, apical segments darker; head, thorax and gaster 
chestnut-brown; legs orange-yellow. 
Cf unknown. 

Material examined 

Holotype §, Zaire: Massif Ruwenzori, Kalonge, 2010 m, Riv. Kamashoro aff. Butahu, 10. ii. 1953 
(Vanschuytbroeck) (MRAC). 

Paratypes. Zaire: 2 $ (BMNH). 

Remarks. Distinguished from prodicus by the relative lengths and shape of the antennal segments. The 
colour pattern is also different. 

Angustacorpa prodicus sp. n. 

(Fig. 8) 

Description. $ antenna short, with segment 3 longer than 4+5, 4 longer than 5, segments 6-13 with 
rhinaria, forming a distinct club (Fig. 8). Head viewed laterally weakly wedge-shaped, with long scattered 



176 J. QUINLAN 

setae; malar grooves percurrent; mandibles tridentate (cf. Fig. 1); occipital carina distinct on lower 
margins; ocelli widely spaced in dorsal view (cf. Fig. 1). Pronotal plate viewed dorsally with a broad medial 
bridge (cf. Fig. 11). [Mesoscutum of holotype damaged by pin.] Scutellar fovea kidney-shaped; scutellar 
disc finely rugose, conical apically, extending past propodeum; scutellar cup long and broad, lateral 
margins with small pits or foveae (cf. Fig. 4). Mesopleural suture fine; metapleural anteroventral cavity 
weakly pubescent; propodeal sculpture with long hairs intermingled. Gaster segment 1 not visible in dorsal 
or lateral views, obscured by ring of pubescence at base of tergite 2, polished; apical third of segment 2 and 
visible parts of 3-5 finely punctate (cf. Fig. 5). Radial cell of fore wing appearing closed. Legs short and 
stout; coxa and femur without conspicuous pubescence. Colour: antenna brownish yellow basally, apical 
segments pale yellow; head and thorax dark chestnut brown; gaster orange-brown; legs yellow. 
Cf unknown. 

Material examined 
Holotype $, Zaire: Kivu, Rutshuru (riv. Fuku), 1250 m, 5.vii.l935 (de Witte) (MRAC). 

Remarks. Closely related to persa (q.v.). 

Angustacorpa triton sp. n. 

(Figs 7, 12, 13) 

Description. $ antenna with segment 3 shorter than 4+5, 4-6 subequal in length, segments 7-13 with 
rhinaria, forming a club (Fig. 7). Head laterally with long scattered setae; malar grooves percurrent; 
mandibles tridentate, produced forward in lateral view (cf. Fig. 1); occipital carina distinct on lower 
margins; ocelli widely spaced in dorsal view. Pronotal plate viewed dorsally with a broad medial bridge (cf . 
Fig. 11); scutellar fovea kidney-shaped; scutellar disc conical apically, extending past propodeum; surface 
of scutellar cup with broken striae. Mesopleural suture weak; metapleuron with aberrant ridges indicated, 
anteroventral cavity without pubescence. Gaster segment 1 short, as broad as long, ridged; tergite 2 with a 
basal hairy ring; segments 2-5 visible in lateral view; apical third of segment 2 and visible parts of 3-5 with 
strong punctures (Fig. 12). Fore wing with radial cell open on margin (Fig. 13). Legs stout; coxa and femur 
without conspicuous pubescence. Colour: antenna brownish yellow; head blackish; thorax and gaster dark 
chestnut brown; legs brownish yellow. 
Cf unknown. 

Material examined 
Holotype $ , Zaire: Mont Hoyo, 1280 m, 7-15.vii.1955 (Vanschuytbroeck) (MRAC). 

Remarks. Distinguished from other species by the 7-segmented antennal club and the strongly punctate 
gaster. 

BOTHROCHACIS Cameron 

Bothrochacis Cameron, 1904: 164. Type species: Bothrochacis erythropoda Cameron, by original designa- 
tion. 

Stirencoela Cameron, 1910: 180. Type species: Stirencoela striaticollis Cameron, by monotypy. [Synony- 
mised by Weld, 1930:139.] 

Ditrupaspis Kieffer, 1910a: 18. Type species: Ditrupaspis semirufa Kiefer, by monotypy. [Synonymised by 
Weld, 1930: 139.] 

Salpictes Kieffer, 1913: 31. Type species Salpictes rufiventris Kieffer, by monotypy. [Synonymised by 
Weld, 1931:223.] 

Diagnosis. $ antenna 13-segmented, clavate, segments 7-13 with rhinaria, forming a distinct club 
(Fig. 15). cf antenna 15-segmented, filiform, segments 3-14 with rhinaria. Head viewed frontally with 
eyes, measured medially, further apart than the height of an eye, frontal area with radiating striae, weakly 
pubescent; malar grooves percurrent, striated on either side; back of head strongly sculptured. Pronotal 
plate strongly produced forwards, posterior and anterior parts separated by a medial bridge with an open 
fovea on each side, lateral margins densely pubescent. Pronotum on either side of pronotal plate crenulate 
or striate. Mesoscutum smooth and polished with a few sparse hairs; lateral bars of scutellum with 
longitudinal striations; scutellar fovea large, smooth and polished, separated by a narrow septum 
extending from the scutellar cup; scutellar cup pear-shaped, apical fovea declined, not visible in dorsal 
view; scutellar disc reticulate-rugose, rounded apically (in some males it can be bicuspid) (Fig. 16). 
Mesopleuron smooth and polished, mesopleural suture complete; metapleuron weakly ridged, anteroven- 



AFROTROPICAL EUCOILIDAE 177 

tral cavity pubescent; propodeal carinae parallel sided though U-shaped. Segment 1 of gaster obscured by 
dense hairy ring (complete on dorsal surface): segment 2 the largest in lateral view, segments 3 and 4 
partially visible; gaster partially or wholly impunctate; hypopygium weakly protruding (Fig. 18). Legs with 
coxae bulbous; femora, tibiae and tarsi pubescent. Fore wing with radial cell open or closed on margin but 
only weakly pigmented; R { thinner than either Rs { or Rs 2 ; surface of wing dotted with hair bases (Fig. 19), 
apical margins of hind wing with a weak hair fringe; wings strongly infuscate basally. Colour: head and 
thorax blackish; gaster bright orange-yellow to black; legs brown-yellow. 

Distribution. A small genus at present known only from Africa. 

Discussion. Bothrochacis is separated from the closely related genus Eucoila. on the unusual declined 
apical fovea on the scutellar cup. Nordlander (1982) places Bothrochacis in his Trybliographa group of 
genera. Five nominal species have been covered in my examination of the genus and three of these I 
consider to be synonymous with the type species B. erythropoda, confirming some of Weld's synonymy. 

Key to species of Bothrochacis Cameron 

Females 

1 Segment 3 of antenna longer than 4+5 (Fig. 31); scutellar disc rugose. 

Antennal segment 5 longer and broader than 4, 6 broader than 5; gaster punctate, black 
(Fig. 35) runVenfris (Kieffer) (p. 178) 

- Segment 3 of antenna subequal to or shorter than 4+5; scutellar disc reticulate-rugose 2 

2 Segment 3 of antenna shorter than 4+5 (Fig. 26); gaster reddish brown; hind tibia with a 

longitudinal ridge (Fig. 28) serratepilosa Benoit (p. 178) 

- Segment 3 of antenna as long as 4+5 (Fig. 15); gaster bright orange-red; hind tibia without a 

longitudinal ridge erythropoda Cameron (p. 177) 

Males 

B. erythropoda is the only species in which the male is known. 

Bothrochacis erythropoda Cameron 
(Figs 15-17, 19) 

Bothrochacis erythropoda Cameron, 1904: 164. LECTOTYPE cf , South Africa (BMNH), here desig- 
nated [examined] [Type no. 7-34]. 

Stirenocoela striaticollis Cameron, 1910: 180. Holotype (J , South Africa (BMNH) [examined]. [Synony- 
mised by Weld 1930: 139.] 

Ditrupaspis semirufa Kieffer, 1910a: 18. Holotype cf, Malawi (MNHU) [examined]. Syn. n. 

Bothrochacis stercoraria Bridwell, 1919: 178. Holotype $, South Africa: Capetown vicinity, iv.1915 
(Bridwelt) (SAM) [not examined]. Syn. n. 

Description. $ antenna with segments 7-13 forming a distinct club, each segment slightly longer than 
wide, segment 3 as long as 4+5, 4-6 subquadrate; cf antenna with segments 3 and 4 swollen, segment 3 
shorter than 4, 4 shorter than 5, 5 and 6 subequal in length, 6-9 subequal but tapering, 9-13 each shorter 
and slender (in lectotype segment 14 and whole of right antenna missing). Face smooth and polished 
medially with weak radiating striae on inner orbits; occiput finely and closely striated. Pronotal plate 
(Fig. 17); pronotum on either side of pronotal plate striate. Mesoscutum without trace of notauli; lateral 
bars of scutellum very finely sculptured dorsally. Metapleuron polished in upper region, sculptured on 
lower margin; lateral margins of propodeum obscured by pubescence; propodeal carinae bowed. Segment 
1 of gaster in form of a narrow ring, weakly crenulate ; apex of tergite 2 and whole of visible part of segment 

3 punctate. Wings without pubescence, margins (except lower margin of hind wing (Fig. 19)) without a 
fringe of hairs ; radial cell of fore wing appears partially open on wing margin due to a lack of pigmentation ; 
cubitus (vein M) indicated, weakly pigmented basally; Rs + M indicated; basal infuscation of wing 
brownish. Legs robust; fore tibia with sharply curved bifid spur; mid and hind tibiae with two single spurs. 
Colour: antenna segment 1 black, remainder reddish brown; gaster orange-brown. 

Material examined 

Bothrochacis erythropoda Cameron, lectotype cf, South Africa: Cape Colony, Grahamstown, ?xi {Daly 
& Sale) (BMNH). Stirenocoela striaticollis Cameron, holotype cf , South Africa: Cape Colony (BMNH). 
Ditrupaspis semirufa Kieffer, holotype cf , Malawi: 'N. Nyasa' (MNHU). 

Kenya: 2 $ (BMNH). South Africa: 26 cf , 9 $ (BMNH); 6 $ (AM). Zaire: 6 2 (MR AC). 



178 J. QUINLAN 

Remarks. One of two male syntypes of B. erythropoda in the BMNH collection is designated as lectotype. 
It bears Cameron's data label 'Cape Colony'. Both specimens, however, bear the name erythropus and not 
erythropoda as published. Weld (1930) found that Stirenocoela striaticollis is synonymous with erythropoda 
and this has been confirmed. I have examined what remains (the antenna, gaster and parts of the legs are 
missing) of the holotype of Ditrupaspis semiruf a. It bears labels 'N. Nyasa, Lamgenburg . . .98, Fulleborn, 
S', 'Co type' [printed], and determination and syntype labels of G. Nordlander. Like Weld (1952) I 
consider it to be conspecific with B. erythropoda. Bridwell (1919) based his description of B. stercoraria on 
10 $ and 2 d" from cow dung or bred from the puparia of Musca lusuria and Lasiopyrellia cyanea 
(Bridwell). He stated that he had been inclined to identify his material as erythropoda except that species 
was said to have the second abscissa of the radius roundly curved. I have not seen specimens of B. 
stercoraria but regard the distinguishing character as too variable to reliably separate species. On the basis 
of the description I therefore synonymise it with B. erythropoda. 

Bothrochacis rufiventris (Kieffer) 

(Figs 31, 34, 35) 

Salpictes rufiventris Kieffer, 1913: 31. Holotype $, Kenya (MNHN) [examined]. 

Description. $ antenna with segment 3 longer than 4+5, 5 longer and broader than 4, 6 broader than 5 
(Fig. 31), 7-13 with rhinaria, each progressively broader and longer than the preceding segment, forming a 
distinct club. Face smooth and polished with radiating striae on inner margins of eyes; occiput with strong 
vertical striations. Mesoscutum with weak anterior parallel lines present, notauli weakly indicated; 
scutellum (Fig. 34); scutellar disc coarsely rugose. Propodeal carinae obscured by dense pubescence on 
lateral margins. Segment 1 of gaster in form of a crenulate ring; parts of segments 3 and 4 punctate 
(Fig. 35). Fore wing with radial cell open on margin; cubitus (R{) indicated; vein Rs + M extending to base 
of Rs and M; hind wing with a fringe of hairs on lower margin. Legs robust, pubescent; fore tibia with 
sharply curved bifid spur; mid and hind tibiae with two single spurs. Colour: antenna reddish brown; gaster 
black. 

Cf unknown. 

Material examined 

Holotype 2, Kenya: British East Africa, 'Afrique Orientale Anglaise', Kajabe, 2100 m, St. no. 27, 
xii.l911(MNHN). 

Remarks. I have seen only the holotype of this species. It is separated from erythropoda by antennal and 
colour characters. 

Bothrochacis serratepilosa Benoit 

(Figs 26-30, 32, 33) 

Bothrochacis serratepilosa Benoit, 1956: 534. Holotype $, Rwanda (MRAC) [examined]. 

Description. $ antenna subclavate, segments 7-13 with rhinaria, segment 3 shorter than 4+5,4 and 5 
subequal in length, 6 and 7 subequal, 7-13 swollen medially, 13 longer than 12 (Fig. 26). Head in lateral 
view wedge-shaped; frontal radiating striae weak, occiput with weak vertical striations. Pronotum on 
either side of pronotal plate strongly striate; pronotal plate with lateral foveae elongate, medial bridge 
narrow (Fig. 27). Scutellar disc truncate apically. Propodeal carinae bowed medially, pubescent on the 
outer margins. Gaster with apex of segment 2 and visible parts of 3-5 punctate; hypopygium pronounced. 
Hind tibia with a distinctive longitudinal ridge (Fig. 28). Fore wing with radial cell open on wing margin 
(Fig. 29), margin weakly pigmented; cubitus (M) not extending to apex of wing; apical and hind margins of 
hind wing with a hair fringe; vein Rs + M not visible but outlined by a brownish hue extending across the 
basal half of the wing; vein Sc + R t with a distinct break (Fig. 29). Colour: entirely reddish brown. 
Cf unknown. 

Material examined 
Holotype $, Rwanda: Terr. Ruhengeri, Kagogo, 1900 m, 29. i. 1953 (Basilewsky) (MRAC). 

Remarks. Distinguished by antennal and colour characters. 



AFROTROPICAL EUCOILIDAE 179 

DIGLYPHOSEMA Foerster 

Diglyphosema Foerster, 1869: 342. Type species: Diglyphosema eupatorii Foerster, by original designation 

and monotypy. 
Amphiglyphosema Benoit, 1956: 546. Type species Amphiglyphosema latesulcatum Benoit, by original 

designation and monotypy. Syn. n. 

Diagnosis. $ antenna 13-segmented, weakly subclavate; cf antenna 15-segmented with modified third 
segment (Fig. 36). Head viewed frontally with malar grooves present with weak striations on either side; 
eyes as far or slightly further apart than the height of an eye. Pronotum with crenulate sculpture viewed 
dorsally; pubescent on either side of pronotal plate which is broad (Fig. 44). Mesoscutum with distinct but 
broken notauli with sparse hairs on apical half of each notaulus which is widened ; scutellum with a large cup 
overhanging scutellar disc which is rugose to punctate; scutellar cup with a large central fovea or depression 
with a ring of smaller foveae close to border of cup. Mesopleural suture distinct with weak striations on 
dorsal margin. Gaster with segment 1 in form of a collar with weak crenulations, this segment sometimes 
obscured by segment 2 which occupies whole of remaining visible part of gaster ; segment 2 of gaster smooth 
and shining, sometimes with a trace of punctures. Radial cell of fore wing open or closed (this is difficult to 
ascertain because the pigmentation is weak); wing surfaces ciliate, margin of wings with a hair fringe; 
cubitus distinct, not always pigmented. Legs generally clear yellow. 

Distribution. Europe, Africa, the New World. 

Discussion. Nordlander (1976) considers the Foerster genera Disorygma, Microstilba, Gronotoma 
and Diglyphosema to constitute a distinct genus-group. These genera with Zaeucoila constitute the 
Gronotoma-group of Nordlander (1982). Benoit (1956) described a further genus Amphiglyphosema 
which he indicated was similar to Diglyphosema. Weld (unpublished notes) suggested that the two genera 
differed in the radial cell. However, I find nothing to justify retaining Amphiglyphosema as a distinct genus 
at the present time. I also consider Eucoilidea Ashmead to belong to the Gronotoma-group. Weld (1952) 
referred to Hedicke's opinion (1930) that Eucoilidea was a synonym of Gronotoma. I have not seen the 
type species of Gronotoma but regard Eucoilidea as distinct (Quinlan, 1986). More extensive study of the 
Gronotoma-group is needed before the generic classification can be further refined. 

Little is known of the host associations of Diglyphosema. Quinlan (1978) cites Melanagromyza 
aeniventris (Fallen), M. nibletti Spencer, M. dettmeri Hering, M. eriolepidus Spencer and M. tripolii 
Spencer as hosts of the European species D. conjungens Kieffer. The only host data for African species 
relate to D. latesulcatum (Benoit), from Melanagromyza phascoli Tryon and Anthomyia centrosematris de 
Meijere. Both of these Diptera are stem borers recorded throughout the Old World tropics. 

Key to the type species and Afrotropical species of Diglyphosema Foerster 

Females 

1 Radial cell of fore wing open on wing margin (Fig. 38); antennal segments 3-9 subequal in 

length; gaster punctate (cf . Fig. 33) eupatorii Foerster (p. 179) 

- Radial cell closed (Fig. 41) ; antennal segments 3-12 each progressively shorter (Fig. 43) ; gaster 

punctate or impunctate 2 

2 Antennal segment 3 shorter than 4; radial cell of fore wing 2-5 times as long as wide; scutellar cup 

as broad apically as basally (Fig. 40); notauli narrowly separated apically (Fig. 40); gaster 
punctate apically (Fig. 39) utica sp. n. (p. 180) 

- Antennal segment 3 subequal to 4; radial cell of fore wing 2-0 times as long as wide (Fig. 42); 

scutellar cup as long as wide, dew-drop shaped (Fig. 48); notauli as far apart apically as basal 
width of a notaulus (Fig. 48) ; gaster impunctate latesulcatum (Benoit) (p. 180) 

Males 

No males of the Afrotropical species are known. 

Diglyphosema eupatorii Foerster 

(Figs 38, 44-46) 

Diglyphosema eupatorii Foerster, 1869: 345. Holotype $, Germany (MNHU) [examined]. 

Description. £ antenna very weakly subclavate, segments 3-9 subequal, segments 10-12 subequal, each 
fractionally broader than preceding one. Head viewed frontally with inner margins of eyes almost parallel; 
malar grooves obscured by striations on either side; face with scattered pubescence. Pronotum either side 



180 J. QUINLAN 

of pronotal plate canaliculate. Mesoscutum with deep notauli converging, but not merging, posteriorly, 
where they are broader and sculptured; base of scutellum with two deep foveae; disc on lateral margins 
punctate; cup large and extending past apex of scutellum, with a large central fovea. Carinae of propodeum 
parallel; mesopleural suture complete, area below suture smooth and shining. Segment 1 of gaster short, 
broader than long, crenulate; segment 2 the only segment visible in lateral view; apical third of gaster 
punctate. Legs orange-yellow. Wings with hair fringe entire; radial cell of fore wing longer than wide, open 
on margin; cubitus not visible (Fig. 38). 
Material examined 

Holotype $, Germany: Aachen (MNHU). 

Remarks. This, the type species of the genus, does not occur in Africa; it is included to facilitate 
comparison with the Afro tropical species. 

Diglyphosema latesulcatum (Benoit) comb. n. 

(Figs 36, 37, 42, 47-49) 

Amphiglyphosema latesulcatum Benoit, 1956: 546. Holotype $, Rwanda (MRAC) [examined]. 

Description. $ antenna subclavate, segments 3 and 4 equal in length; cf antenna (Fig. 36) with segments 
4-15 moniliform. Face with long scattered pubescence; malar space with distinct striations; frontal line 
raised. Pronotal plate large, not protruded, with two foveae medially. Mesoscutum with notauli converg- 
ing strongly half way towards scutellum, the apical third broad and sculptured, in the form of a series of 
linked foveae (Fig. 48); scutellar disc viewed dorsally visible only on the lateral margins, surface of disc 
reticulate-rugose with long scattered hairs forming a fringe apically; scutellum with two foveae at base. 
Carinae of propodeum converging basally, parallel medially; mesopleuron smooth and polished. Segment 
1 of gaster usually obscured by segment 2; segment 1 short, broader than long, crenulate; segment 2 with a 
few scattered hairs at base; gaster impunctate. Radial cell of fore wing closed; base of wings fumate; Rs + 
M and M not pigmented or indicated. Colour: antenna light brown basally, becoming darker towards apex; 
head and thorax black; gaster chestnut brown; legs orange-yellow. 

Material examined 

Holotype $ , Rwanda: Ruhengerri, Kagogo, 1900 m, 29.L1953 (Basilewsky) (MRAC). 

Remarks. Differs from the European species of Diglyphosema mainly in having the radial cell closed and 
the gaster not densely punctate and by the shape of the scutellar cup. A long series of males, possibly of this 
species, has been examined but there is not sufficient evidence to justify a positive association. 

Diglyphosema utica sp. n. 

(Figs 39, 40, 41, 43) 

Description. $ antenna subclavate, segment 3 shorter than 4, 4 longer than 5, 5 and 6 subequal in length, 
7-13 each slightly, progressively shorter (Fig. 43). Inner margins of eyes with long scattered pubescence; 
malar space with distinct striations on either side; frontal line raised. Pronotal plate, not protruded, viewed 
frontally posterior part longer than anterior part, foveae either side of medial bridge open (Fig. 44). 
Mesoscutum smooth and polished, with notauli converging sharply two-thirds towards scutellum, 
separated basally by a narrow stem, wine glass-shaped (Fig. 40); lateral bars of scutellum smooth and 
polished, short and broad; scutellar foveae shallow, polished; scutellar disc reticulate-rugose on lateral 
margins, its apex obscured by scutellar cup; scutellar cup longer than broad, as broad apically as basally, 
the ring of foveae extending from apex of rim to two-thirds of way towards scutellar fovea. (Fig. 40); apex 
of disc with long hairs forming a fringe. Propodeum obscured by dense pubescence; mesopleuron smooth 
and polished; mesopleural suture arched; metapleuron and side of propodeum densely pubescent. 
Segment 1 of gaster in form of a crenulate ring; segment 2 finely punctate apically. Radial cell of fore wing 
closed on margin (Fig. 41); base of wings fumate; Rs + M and M not indicated. Colour: antenna yellowish 
basally, becoming darker towards apex; head and thorax black; gaster chestnut brown; legs orange-yellow. 
Cf unknown. 

Material examined 

Holotype ?, Nigeria: W. State, Ile-Ife, v.1973 (Medler) (BMNH). 
Paratypes. Ivory Coast: 2 $ (BMNH). 

Remarks. Very closely related to latesulcatum but distinguished by antennal, wing and scutellar cup 
characters. 



AFROTROPICAL EUCOILIDAE 181 

EMARGO Weld 

Emargo Weld, 1960: 195. Type species: Emargo eciton Weld, by original designation and monotypy. 

Diagnosis. $ antenna 13-segmented (rarely 12), segment 1 longer than 2, swollen apically, segment 2 
subquadrate, very broad, segments 7-13 in type species forming an indistinct club. Head broader than 
thorax. Pronotal plate protruding, lateral fovea open. Mesoscutum smooth and polished, notauli fine, 
complete; scutellar fovea weakly indicated; lateral bars of scutellum broad basally, short and polished; 
scutellar disc reticulate-coriaceous, rounded apically; scutellar cup large, occupying almost entire dorsal 
surface of scutellum (Fig. 51). Mesopleural suture absent, surface of mesopleuron smooth and polished; 
metapleuron polished, anteroventral cavity hairless; propodeum with dense woolly pubescence laterally. 
Segment 1 of gaster obscured by propodeal pubescence and pubescence at base of tergite 2; tergites 2-6 
visible in lateral view, 2 shorter than 3 (Fig. 23). Wing surfaces pubescent, fore wing incised at apex, distal 
margins with fringe of hairs; radial cell of fore wing very small, open on margin (Fig. 21); hind wings 
narrow. Legs with hind coxa swollen with pubescence basally. 

Distribution. Mexico, Marianas Is, Africa, Madagascar, Sulawesi, Australia. 

Discussion. Emargo was described and placed by Weld in the Figitinae. He stated that it differed from all 
other Figitinae in having the fore wings emarginate. After studying the genus I conclude that it belongs to 
the Eucoilidae, to which it is transferred. Contrary to Weld's description of the scutellum, the cup is 
exceptionally large and occupies the whole of the dorsal surface. Also it has a typical eucoilid pronotal 
plate, dense pubescence on the lateral margins of the propodeum and a hairy ring at the base of tergite 2. 
However, unlike the majority of eucoilids, segment 2 of the gaster is not the largest. I consider the 
relatively small size of segment 2 to be a primitive character. Apart from the type species and the 
Afrotropical species dealt with here I have seen specimens of Emargo from Sulawesi and Australia. The 
scutellar cup in this latter material is weakly represented on a polished scutellar disc. Two males and one 
female from Australia belong to a species very close to one of the Afrotropical species. 

It is not possible to run this genus to a family in Weld's key (1952). In my key to families and subfamilies 
(Quinlan, 1979) it would run to Eucoilidae but for the relative sizes of gastral segments 2 and 3. Couplet 5 
(Quinlan, 1979: 90) should be amended to read 'Segments 2 and 3 of gaster generally fused, without visible 
suture, exceptionally with segment 2 differentiated.' 'The differentiation of segment 2 in Emargo is similar to 
the condition found in Figitinae. The weakly delineated scutellar cup (cf. Fig. 59) can also appear very 
similar to the smooth scutellum of figitines. 

The only two males known from Africa have 14-segmented antennae. Emargo has in common with 
Glauraspidia, Pseudopsichacra and the new genus Sirenes a densely woolly pronotal plate, densely 
pubescent propodeal and basal gastral segments, and no mesopleural suture. 

Weld (1960) stated that Emargo eciton was taken by Berlese funnel extraction from refuse deposits of the 
army ant Eciton burchelli (Westwood) and presumed its host to be one or more flies of the families 
Phoridae, Muscidae or Sarcophagidae. 

Key to the Afrotropical species of Emargo Weld 

Females 

1 Antenna with 7-segmented club, each segment a little longer than broad (Fig. 53). 

Legs very pale yellow laverna sp. n. (p. 183) 

- Antennal club 8-10 segmented, each with rhinaria 2 

2 Antennal club 8-segmented (Figs 54, 56) 3 

- Antennal club 9- 10-segmented (Figs 55 , 57) 4 

3 Antennal club with conspicuous rhinaria, each segment 3 times as long as broad (Fig. 54) ; apex 

of wing broad, weakly incised pexus sp. n. (p. 185) 

- Antennal club with inconspicuous rhinaria, each segment less than 3 times as long as broad 

(Fig. 56); apex of wing narrow, strongly incised cantus sp. n. (p. 182) 

- Antennal club 9-segmented 5 

- Antennal club 10-segmented 9 

5 Antenna 12-segmented, segment 3 minute (Fig. 55), shorter than segment 4. 

Antennal club segments sharply indicated, each with strong rhinaria and conspicuous 
setae (Fig. 55); apex of wing sharply incised, broad capito sp. n. (p. 183) 

- Antenna 13-segmented, segment 3 equal in length to 4 or longer 6 

6 Antennal segments 3 and 4 equal in length , club segments 3 times as long as broad (indicated by 

strong rhinaria) (Fig. 57). 

Apex of wings strongly incised ascia sp. n . (p. 182) 



182 J. QUINLAN 

- Antennal segment 3 longer than 4 , apical segments variable in proportions 7 

7 Antennal club segments 2 times as long as broad (blackish apically, rhinaria present) (Fig. 58). 

Wings narrow apically, sharply incised matius sp. n. (p. 183) 

- Antennal club segments not less than 3 times as long as broad 8 

8 Apical segments of antenna very pale; lateral bars of scutellum polished, notauli indistinct. 

Scutellar cup long and narrow (Fig. 59) palloris sp. n. (p. 185) 

- Apical segments of antenna brownish black; lateral bars of scutellum sculptured, notauli 

distinct (Fig. 60) numa sp. n. (p. 184) 

9 Antennal segment 3 shorter than 4. 

Antennal club segments 3 times as long as broad (Fig. 61) themis sp. n. (p. 186) 

- Antennal segment 3 as long as or longer than 4 10 

10 Antennal segment 3 distinctly longer than 4. 

Antennal club segments 3 times as long as broad, with rhinaria (Fig. 62). vacuna sp. n. (p. 186) 

- Antennal segments 3 and 4 equal in length 11 

11 Club segments of antenna 3 times as long as broad (Fig. 63), rhinaria inconspicuous. 

micipsa sp. n. (p. 184) 

- Club segments of antenna slightly longer than broad, rhinaria conspicuous (Fig. 64). 

peleus sp. n. (p. 185) 

Males 

The only two male specimens from Africa cannot be associated with any particular species. 



Emargo ascia sp. n. 

(Figs 51, 52, 57, 65) 

Description. 5 antenna with segments 3 and 4 subequal in length, 2 wider than 3 or 4, 5-13 with rhinaria 
forming a club (Fig. 57). Face smooth and polished with scattered hairs; malar grooves not indicated; 
anterior tentorial pits distinct; eyes large. Pronotal plate projected forward, medial bridge broader than 
the open lateral fovea; pronotum with long scattered setae. Notauli not percurrent, with a few scattered 
setae in their place; lateral bars of scutellum small triangular; scutellar fovea shallow; scutellar disc smooth 
and polished; scutellar cup long and narrow, weakly defined; apex of scutellar disc round, surface weakly 
sculptured (Fig. 51). Propodeum obscured laterally and dorsally by dense woolly pubescence; area below 
ventral region of mesopleuron pubescent. Gaster with tergite 3 the largest, tergites 4-7 partially visible 
(Fig. 52) ; hypopygium small. Wing surfaces densely pubescent, margins with a long apical hair fringe; veins 
thick (Fig. 65). Colour: antenna yellowish brown; head brownish; thorax and gaster brownish; legs pale 
yellow. 

Cf unknown. 

Material examined 
Holotype $, Madagascar: Beresty 12 km, NW. Ambossary, 5-15. v. 1983 (Noyes & Day) (BMNH). 
Paratypes. Madagascar: 2 $ (BMNH). Zambia: 1 $ (BMNH). 

Remarks. Distinguished from closely related species with a 9-segmented antennal club by the proportions 
of the antennal segments. 

Emargo cantus sp. n. 

(Figs 56, 66, 67) 

Description. $ antenna weakly subclavate, segment 2 subequal in length to 3 but thicker, 3 longer than 4, 4 
and 5 subequal in length, 6-13 forming a weak club, rhinaria weakly indicated (Fig. 56). Face smooth and 
shining with sparse scattered hairs medially; malar grooves not indicated; anterior tentorial pits pro- 
nounced. Pronotal plate projected forward, medial bridge narrow between the open lateral fovea. Notauli 
percurrent, with 2 or 3 long setae present; scutellar fovea polished, semi-circular; scutellar disc smooth and 
polished; scutellar cup aberrant, faint in dorsal view, long and narrow, declined (Fig. 66). Propodeum 
obscured laterally and dorsally by dense woolly pubescence; ventral margin of mesopleuron pubescent. 
Gaster with tergite 3 the largest, tergites 4-7 partially visible (cf. Fig. 52); hypopygium not prominent. 
Wing surfaces densely pubescent, margins with a long apical hair fringe; veins thick (Fig. 67). Colour: 
antenna, head, thorax and gaster orangy-brown; legs pale yellow. 
Cf unknown. 



AFROTROPICAL EUCOILIDAE 183 

Material examined 

Holotype $, Zimbabwe: Salisbury, Chishawasha, i. 1979 (Watsham) (BMNH). 
Paratypes. Zimbabwe: 3 ? (BMNH). Zaire: 1 $ (MRAC). 

Remarks. This species is most closely related topexus (q.v.). 

Emargo capito sp. n. 

(Figs 55, 68) 

Description. $ antenna 12-segmented, segment 3 minute, 4-12 with faint rhinaria, forming a distinct club 
(Fig. 55). Face smooth and shining, with scattered hairs; malar space small, grooves not indicated; eyes 
large, further apart than the height of an eye; head viewed dorsally large, almost square, as wide as thorax. 
Pronotal plate projected forward, medial bridge broad, the lateral foveae small, with dense pubescence; 
pronotum with a few sparse hairs. Notauli aberrant, with a few long setae in their place; lateral bars of 
scutellum triangular; scutellar fovea shallow; scutellar disc smooth and polished; scutellar cup weakly 
indicated, a weak apical fovea indicated (Fig. 68). Propodeum obscured laterally and dorsally by dense 
woolly pubescence. Gaster with tergite 3 the largest, tergites 4-7 partially visible (cf. Fig. 52); hypopygium 
pronounced. Apical margins of wings with a long hair fringe; radial cell of fore wing minute; apex of fore 
wings excised (cf. Fig. 21). Colour: antenna light brown; head dark brown; thorax orange-yellow; gaster 
light brown; legs pale yellow. 
Cf unknown. 

Material examined 
Holotype $, Madagascar: Tarn Perinet, 27.iv.1983 (Noyes & Day) (BMNH). 

Remarks. This species is exceptional among the Afrotropical species of the genus in having 12-segmented 
antennae with the third segment very reduced. Two males and one female from Australia are very close to 
this species but until further material is seen or the Australian cynipoid fauna is examined no decision can 
be made on their status. 

Emargo layer na sp. n. 

(Fig. 53) 

Description. $ antenna with segment 2 longer and wider than 3, 3 longer than 4, 4 and 5 subequal in 
length, 6 shorter than 5, 7-13 as wide as long forming a distinct club (Fig. 53). Face smooth and polished 
with scattered hairs; malar grooves aberrant; eyes small, almost round. Pronotal plate projected forward, 
medial bridge as wide as the open lateral fovea; pronotum polished, without hairs. Notauli weakly 
indicated, with a few scattered hairs in their place; lateral bars of scutellum small triangular; scutellar 
fovea shallow; scutellar disc smooth and polished; scutellar cup long and narrow, very weakly defined 
(cf. Fig. 66). Propodeum obscured laterally and dorsally by dense woolly pubescence. Gaster with tergite 3 
the largest, tergites 4-7 partially visible; hypopygium small. Wing surfaces densely pubescent, margins 
with a long apical hair fringe; apex of fore wing strongly incised; veins thick (cf. Fig. 67). Colour: antenna 
yellowish brown; head brownish; thorax and gaster yellowish brown; legs pale yellow. 
Cf unknown. 

Material examined 

Holotype $, Zimbabwe: Salisbury, Chishawasha, ii.1979 (Watsham) (BMNH). 
Paratype. Kenya: 1 $ (BMNH). 

Remarks. This is the only species of the genus with a 7-segmented antennal club. 

Emargo matius sp. n. 

(Figs 58, 69) 

Description. $ antenna with segment 2 subequal in length to 3 but wider, 4 shorter than 3, 5-13 with 
rhinaria forming a distinct club (Fig. 58). Face smooth and polished with scattered hairs medially around 
clypeal region; malar grooves weakly indicated; head viewed dorsally almost as wide as long. Pronotal 
plate projected forward, medial bridge wide, the lateral foveae open; pronotum with long scattered setae. 
Notauli percurrent, with long setae obscuring them; lateral bars of scutellum small; scutellar fovea small 
and shallow; scutellar disc smooth and polished with long setae; scutellar cup long and narrow, weakly 
indicated (Fig. 69). Propodeum obscured laterally and dorsally by dense woolly pubescence; area below 



184 J. QUINLAN 

ventral region of mesopleuron densely pubescent. Gaster with tergite 3 the largest, tergites 4-7 partially 
visible (cf. Fig. 23); hypopygium small. Wing surfaces densely pubescent, margins with a long apical hair 
fringe; apex of fore wing strongly incised; veins thick (cf. Fig. 67). Colour: antenna yellowish brown; head, 
thorax and gaster brownish; legs yellow. 
O" unknown. 

Material examined 

Holotype ?, Zimbabwe: Chishawasha, i.1979 (Watsham) (BMNH). 

Paratypes. Zimbabwe: 3 $ (BMNH). Cameroun: 1 $ (BMNH). Kenya: 1 $ (BMNH). Zaire: 1 $ 
(BMNH). 

Remarks. Distinguished from ascia, palloris and numa on the shape and proportions of the antennal 
segments, these four species are otherwise but little distinct. 

Emargo micipsa sp. n. 

(Figs 23, 63) 

Description. 5 antenna clavate, with segments 2 and 3 subequal in length, 2 wider than 3, 4-13 subequal in 
length, swollen medially, rhinaria distinct (Fig. 63). Face smooth and polished; malar grooves not 
indicated; anterior tentorial pits distinct; eyes further apart measured medially than the height of an eye. 
Pronotal plate projected forward, medial bridge broad, the lateral foveae open; pronotum with sparse long 
setae. Notauli very weakly indicated in part with long scattered hairs in their place ; lateral bars of scutellum 
triangular; scutellar fovea large semi-circular shallow; scutellar disc smooth and polished; scutellar cup 
long and narrow (cf. Fig. 51). Propodeum obscured laterally and dorsally by dense woolly pubescence; 
area below ventral margin of mesopleuron pubescent. Gaster with tergite 3 the largest, tergites 4-7 
partially visible (Fig. 23); hypopygium not prominent. Wing surfaces densely pubescent, margins with a 
long apical hair fringe; apex of fore wing strongly incised; veins thick (cf. Fig. 21). Colour: antenna 
yellowish; head brownish; thorax and gaster brownish yellow; legs pale yellow. 
Cf unknown. 

Material examined 

Holotype 9, Cameroun: Nkoemvon, 13.vii.— 24.viii.1980 (Jackson) (BMNH). 
Paratypes. Cameroun: 3 $ (BMNH). Madagascar: 2 5 (BMNH). Zaire: 3 $ (MRAC). 

Remarks. One of a group of four species with a 10-segmented antennal club, micipsa can be distinguished 
from the others (peleus, themis and vacuna) by the proportions of the antennal segments and, in common 
with vacuna, micipsa has obsolete notauli. 

Emargo numa sp. n. 

(Figs 60, 70, 71) 

Description. $ antenna weakly subclavate, with segment 2 shorter than 3 but thicker, 3 longer than 4, 4 
shorter than 5, 5-13 with rhinaria forming a club (Fig. 70). Face smooth and polished with scattered hairs; 
malar grooves absent. Pronotal plate small, projected forward, the lateral foveae open, weakly pubescent; 
pronotum with a few long sparse setae. Notauli distinct, percurrent, a few sparse hairs present; lateral bars 
of scutellum sculptured; scutellar fovea shallow; scutellar disc smooth and polished; scutellar fovea 
shallow; scutellar disc smooth and polished; scutellar cup long and narrow, weakly defined; apex with small 
pit or fovea, surface of scutellum with long sparse setae (Fig. 60). Propodeum obscured laterally and 
dorsally by dense woolly pubescence; ventral margin of mesopleuron pubescent. Gaster with tergite 3 the 
largest, tergites 4-7 partially visible (cf. Fig. 52); hypopygium not prominent. Wing surfaces densely 
pubescent, margins with a long apical hair fringe; veins thick (cf . Fig. 21). Colour: antenna brownish; head, 
thorax and gaster orange-brown; legs yellow. 
C? unknown. 

Material examined 
Holotype $, Zimbabwe: Salisbury, Chishawasha, i.1979 (Watsham) (BMNH). 
Paratype. Zimbabwe: 1 $ (BMNH). 

Remarks. Differs from palloris in the lateral bars of the scutellum and shape of the scutellar cup. 



AFROTROPICAL EUCOILIDAE 185 

Emargo pallor is sp. n. 

(Figs 59, 72) 

Description. $ antenna filiform, with segment 2 shorter than 3, 3 longer than 4, 4 and 5 subequal in length, 
5-13 with rhinaria forming a very weak club (Fig. 72). Face smooth and polished with scattered hairs; 
malar grooves weakly indicated; head as broad as thorax. Pronotal plate projected forward, medial bridge 
wide; pronotum with long scattered setae. Notauli aberrant, not percurrent, a line of scattered hairs in their 
place; lateral bars of scutellum small triangular; scutellar fovea shallow; scutellar disc hardly visible; 
scutellar cup long and narrow, only weakly indicated, smooth and polished with a few long scattered setae; 
apex of scutellar disc not visible in dorsal view (Fig. 59). Propodeum obscured laterally and dorsally by 
dense woolly pubescence; area below ventral margin of mesopleuron pubescent. Gaster with tergite 3 the 
largest, tergites 4-7 partially visible (cf. Fig. 23); hypopygium pronounced. Wing surfaces densely 
pubescent, margins with a long apical hair fringe; apex of fore wing strongly incised; radial cell short and 
thick (cf. Fig. 67). Colour: antenna yellow with apical segments pale, almost white; head, thorax and gaster 
yellowish brown; legs yellowish. 
Cf unknown. 

Material examined 

Holotype $, Zaire: Massif Ruwenzori, riv. Kamusmge, 1700 m, a.f. Ndama, 3.xi.l953 {Vanschuyt- 
broeck & Hendrrickx) 6143-45 (MR AC). 

Paratypes. Zaire: 3 $ (MRAC, BMNH). 

Remarks. Separated from numa by the lateral bars of scutellum and the scutellar cup. 

Emargo pe/eus sp. n. 

(Figs 64, 73) 

Description. $ antenna clavate, with segment 2 longer and wider than 3, 3 and 4 subequal in length but 
thinner, 4-13 a little longer than wide with rhinaria, forming a club (Fig. 64). Face smooth and polished 
with sparse scattered hairs; malar grooves absent; head as broad as thorax. Pronotal plate projected 
forward, the open lateral foveae small, densely pubescent; pronotum with long scattered setae. Notauli 
distinct, percurrent, a few long hairs present; lateral bars of scutellum small triangular; scutellar fovea 
shallow; scutellar disc smooth and polished, square apically ; scutellar cup long and narrow, extending past 
apex of scutellum (cf. Fig. 66). Propodeum obscured laterally and dorsally by dense woolly pubescence; 
ventral margin of mesopleuron densely pubescent. Gaster with tergite 3 the largest, remaining tergites 
partially visible; hypopygium short and broad (Fig. 73). Wing surfaces densely pubescent, margins with a 
long apical hair fringe; radial cell small; veins thick (cf. Fig. 65). Colour: antenna yellow; head, thorax and 
gaster orange-brown; legs yellow. 
Cf unknown. 

Material examined 

Holotype $, Cameroun: Nkoemvon, vii.-viii. {Jackson) (BMNH). 
Paratypes. Cameroun: 3 § (BMNH). Madagascar: 1 $ (BMNH). 

Remarks. Most closely related to micipsa (q.v.). 

Emargo pexus sp. n. 

(Figs 54, 74) 

Description. $ antenna filiform, with segment 2 subequal in length to 3 but thicker, 3 longer than 4, 
shorter than 5, 6-13 with pronounced rhinaria forming a club (Fig. 54). Face smooth and polished; malar 
grooves not indicated; anterior tentorial pits distinct; eyes measured medially further apart than the height 
of an eye. Pronotal plate projected forward, medial bridge broad. Notauli percurrent; lateral bars of 
scutellum elongate with weak sculpture on lateral margins; scutellum large, scutellar disc smooth and 
polished, narrow apically; scutellar cup weakly indicated, apex with a fovea (Fig. 74). Propodeum 
obscured laterally and dorsally by dense woolly pubescence; ventral margin of mesopleuron pubescent. 
Gaster with tergite 3 the largest, tergites 4-7 partially visible (cf. Fig. 23); hypopygium prominent. Wing 
surfaces densely pubescent, margins with a long apical hair fringe; apex of fore wing weakly incised; veins 
thick (cf. Fig. 21). Colour: antenna brownish yellow head, thorax and gaster blackish; legs yellow. 
Cf unknown. 



186 J. QUINLAN 

Material examined 
Holotype $, Madagascar: TamPerinet, 27.iv.-3.v.l983 (Noyes & Day) (BMNH). 
Paratype. Madagascar: 1 5 (BMNH). 

Remarks. This species and cantus are the only Afrotropical species in the genus with an 8-segmented 
antennal club. They can be distinguished by differences in the shape of the wings and in the proportions of 
the antennal segments. Also their distributions are disjunct, though this could be merely a reflection of the 
paucity of collections of microhymenoptera from the Afrotropical region. 

Emargo themissp. n. 

(Fig. 61) 

Description. $ antenna filiform, with segment 2 subequal in length to 3 but wider, 3 thinner and shorter 
than 4, 4-13 with rhinaria indicated, weakly swollen medially (Fig. 61). Face smooth and polished with 
scattered hairs; malar grooves not indicated; anterior tentorial pits distinct; eyes large, further apart 
measured medially than the height of an eye. Pronotal plate projected forward, medial bridge narrow. 
Notauli percurrent, a few sparse hairs present; lateral bars of scutellum triangular; scutellar fovea large, 
shallow; scutellar disc with aberrant sculpture; scutellar cup long and narrow (cf. Fig. 51). Propodeum 
obscured laterally and dorsally by dense woolly pubescence; area below ventral margin of mesopleuron 
densely pubescent. Gaster with tergite 3 the largest, tergites 4-7 partially visible (cf. Fig. 52); hypopygium 
small. Wing surfaces densely pubescent, margins with a long apical hair fringe; apex of fore wing strongly 
incised; veins thick (Fig. 65). Colour: antenna yellowish brown; head, thorax and gaster dark brown; legs 
pale yellow. 
Cf unknown. 

Material examined 
Holotype $, Cameroun: Nkoemvon, 13.vii.— 24.viii.1980 (Jackson) (BMNH). 
Paratypes. Cameroun: 4 $ (BMNH). Madagascar: 2 5 (BMNH). 

Remarks. This apparently widespread species belongs to the group having a 10-segmented antennal club 
(see remarks on micipsa). 

Emargo vacuna sp. n. 

(Fig. 62) 

Description. $ antenna clavate, segment 2 shorter but wider than 3, longer than 4, 4-13 more than twice 
as long as wide, with rhinaria weakly indicated (Fig. 62). Face smooth and polished with scattered setae; 
malar grooves not indicated; anterior tentorial pits distinct; eyes measured medially further apart than the 
height of an eye. Pronotal plate projected forward, posterior part of plate angular, medial bridge broad, 
the lateral foveae with tufts of pubescence; pronotum with sparse hairs. Notauli not percurrent, with a few 
long hairs in their place; lateral bars of scutellum triangular; scutellar fovea large, shallow and polished; 
scutellar disc smooth and polished; scutellar cup long and narrow, weakly outlined with a few long setae 
(cf. Fig. 69). Propodeum obscured laterally and dorsally by dense woolly pubescence; area below ventral 
margin of mesopleuron densely pubescent. Gaster with tergite 3 the largest, tergites 4-7 partially visible 
(cf. Fig. 52); hypopygium small. Wing surfaces densely pubescent, margins with a long apical hair fringe; 
veins thick (Fig. 65). Colour: antenna light brown; head, thorax and gaster orange-brown; legs pale yellow. 
Cf unknown. 

Material examined 
Holotype 5 , South Africa: Pondoland, Port St John, 6-25. ii. 1924 (Turner) (BMNH). 

Remarks. Distinguished from other species by antennal characters (see remarks on micipsa). 

EUCOILA Westwood 

Eucoila Westwood, 1833: 494. Type species: Eucoila crassinervis Westwood, by original designation. 
Psilodora Foerster, 1869: 354. Type species: Cothonaspis boyenii Hartig, by original designation. 

[Synonymisedby Weld, 1952: 212.] 
Lytosema Kieffer, 1901: 159. Type species: Eucoela guerini Dahlbom, by subsequent designation of 

Ashmead, 1903: 67. [Synonymised by Weld, 1931: 222.] 
Psilodoropsis Hedicke, 1913: 413. Type species: Psilodoropsis conradti Hedicke, by original designation 

and monotypy. [Synonymised by Nordlander, 1981: 398.] 



AFROTROPICAL EUCOILIDAE 187 

Diagnosis. $ antenna 13-segmented, clavate, segments 6-13 with rhinaria and forming a distinct club in 
the type species; cf antenna 14-segmented, filiform, segments 3-14 with rhinaria, subequal in length 
(Fig. 78). Head viewed frontally almost square (Fig. 77); inner margins of eyes subparallel, further apart 
measured medially than height of an eye; malar grooves percurrent, striated either side of groove near 
clypeus; clypeus and mandibles with long setae; head polished on occiput and vertex. Pronotal plate 
strongly produced in form of a collar viewed dorsally, medial bridge with a lateral fovea on either side, 
open, with dense pubescence. Mesoscutum smooth and polished, with sparse scattered hairs, notauli not 
present; scutellum with strongly striated lateral bars, scutellar foveae window-like (Nordlander, 1982), 
large deep; scutellar plate (cup) oval, weakly excavate, with a fovea on the apex, visible in dorsal view; 
scutellar disc reticulate-rugose, rounded apically (Fig. 79). Mesopleuron smooth and polished, mesopleu- 
ral suture (carina) complete; sides of pronotum and precoxal area striated; metapleuron strongly ridged, 
anteroventral cavity striated; propodeal carinae angular. Segment 1 of gaster obscured by a dense woolly 
ring of pubescence (complete on dorsal surface); segments 2-4 visible in lateral view, apical quarter of 
segment 2 not visible; parts of 3 and 4 densely punctate; hypopygium with long subapical setae. Coxae 
bulbous, smooth; femora, tibiae and tarsi weakly pubescent. Fore wing with a distinctly closed radial cell in 
type species; vein R { thinner than either Rs x or Rs 2 ; surfaces of wings with dotted hair bases, apical margins 
of wings with weak hair fringe (cf. Fig. 81); wings weakly infuscated. Colour: antenna black basally, club 
yellowish brown; head, thorax and gaster blackish brown; legs brownish yellow. 

Distribution. Europe, Africa, the New World. 

Discussion. Nordlander (1982) regards Eucoila as the sister-group of Trybliographa and Bothrochacls and 
he divides the genus into two species-groups: the nudipennis-group and the crassinerva-group. Eucoila 
differs primarily from Bothrochacis by the form of the scutellum, the apical fovea on the scutellar cup being 
sharply declined in Bothrochacis (Fig. 30) and not visible in dorsal view. Trybliographa always has the 
surface of the wings pubescent and the marginal fringe of hair present (Fig. 81). Both species-groups of 
Eucoila lack pubescence on the surface of the wings, while the nudipennis-group has a reduced subalar pit. 
Trybliographa and the crassinerva-group of Eucoila, together with Leptopilina, have an area behind the 
postero ventral edge of the metapleuron completely free of pubescence. 

Five Afrotropical species have been described in Eucoila, namely, E. cavernicola Kieffer and E. 
kilimandjaroi Kieffer transferred to Pseudeucoila by Weld (1952), E. flagellicornis Kieffer transferred to 
Psichacra by Weld (1952), and E. camerounensis Risbec which cannot be satisfactorily identified (see 
below). Four new species are described and keyed here. 

Key to Afrotropical species of Eucoila Westwood 

Females 

1 Antennal club 7-segmented (Fig. 82) 2 

- Antennal club 8-9 segmented. 

Radial cell of fore wing distinctly open (Fig . 92) ; mesopleural suture distinct 3 

2 Mesopleural suture indistinct, obscured by microsculpture (Fig. 88); radial cell of fore wing 

elongate, almost closed on wing margin (Fig. 90); pronotal plate (Fig. 89); antennal segment 3 
shorter than 4+5 (Fig. 82) marina sp. n. (p. 188) 

- Mesopleural suture distinct, not obscured by sculpture; radial cell of fore wing squarish (cf. 

Fig. 80); pronotal plate (Fig. 86); antennal segment 3 subequal to 4+5 (Fig. 84). 
bantia sp. n. (p. 187) 

3 Antennal club 8-segmented; scutellar disc rounded apically (cf . Fig. 87) veleda sp. n. (p. 189) 

- Antennal club 9-segmented (Fig. 85); scutellar disc bicuspid apically (Fig. 91). erinna sp. n. (p. 188) 

Males 

Only the male of E. marina sp. n. is known. 

Eucoila bantia sp. n. 

(Figs 84, 86, 87) 

Description. $ antenna with segments 7-13 with rhinaria and forming a club (Fig. 84), segment 3 longer 
than 4, 4-6 subequal in length, club segments 2 times as long as wide. Head viewed frontally with eyes as far 
apart as the height of an eye measured medially; face smooth and polished with sparse scattered 
pubescence. Pronotal plate (Fig. 86); pronotum either side of pronotal plate pubescent. Scutellar fovea 
smooth and polished, shallow; scutellar cup with medial area transversely striated; scutellar disc Fig. 87. 
Metapleuron polished; anteroventral cavity pubescent; propodeal carinae weakly bowed, pubescent on 



188 J. QUINLAN 

outer margins; nucha obscured by pubescence. Segment 1 of gaster in form of a narrow ring; segment 2 the 
largest, occupying most of the lateral area; segment 3 partially visible; gaster impunctate; hypopygium 
broad apically. Radial cell of fore wing partially open on wing margin (cf. Fig. 92); lateral and medial area 
of wings infuscate. Coxae longer than wide; hind coxa with distinct comb of hairs on upper posterior 
margin; femora and tibiae long and narrow, femora swollen medially, tibiae swollen basally; metatarsus as 
long as remaining tarsal segments. Colour: antenna brownish yellow; head and thorax black, gaster 
chestnut brown; legs dark orange-brown. 
Cf unknown. 

Material examined 
Holotype $, Zimbabwe: Salisbury, Chishawasha, ii.1978 (Watsham) (BMNH). 
Paratypes. Zimbabwe: 4 $ (BMNH). Botswana: 2 $ (BMNH). Zaire: 7 $ (MRAC). 

Eucoila camerounensis Risbec 

Eucoila camerounensis Risbec, 1956: 160. Syntypes 2 cf, 5 $, Cameroun: Mil. Garoua, vii.1954 
(Decamps) (?lost). 

I have not been able to locate the type material of this species and on the basis of the description and figure 
cannot reach any conclusions about its identity. 

Eucoila erinna sp. n. 

(Figs 83, 85, 91, 92) 

Description. $ antenna with segments 5-13 with rhinaria and forming a club (Fig. 83), segment 3 longer 
than 4, 4 shorter than 5. Head viewed frontally with eyes slightly further apart than the height of an eye 
measured medially; inner orbits of face with reticulate-rugose sculpture converging towards clypeus, 
extending to malar grooves, face and clypeus with scattered setae; occiput strigose. Pronotal plate 
cf. Fig. 86; outer parts angular. Lateral bars of scutellum very weakly striated; scutellar fovea deep, 
polished; scutellar cup almost circular with transverse sculpture; scutellar disc with apex weakly bicuspid 
(Fig. 91). Metapleuron weakly ridged basally; anteroventral cavity pubescent; propodeal carinae weakly 
bowed basally, outer margins of carina and of nucha densely pubescent. Segment 2 of gaster the largest, 
apical quarter finely punctate; segments 3 and 4 partially visible; hypopygium broad apically, with 
sub-basal hairs. Fore wing of female bare, apical half with dotted hair bases; radial cell open; posterior 
angle of wing with a weak fringe of hairs (more prominent on hind wing); medial area of wings infuscate. 
Coxae longer than wide, bulbous, pubescent; trochanters longer than wide; femora swollen basally, long; 
tibiae long and narrow basally; tarsi narrow, metatarsus as long as remaining tarsal segments combined. 
Colour: antenna orange-brown; head and thorax black; gaster and legs orange. 
Cf unknown. 

Material examined 

Holotype $, Zaire: Mabanga, 29.ix.1952 (De Saeger) (MRAC). 
Paratypes. Zaire: 13 $ (MRAC). 

Eucoila marina sp. n. 

(Figs 82, 88-90) 

Description. $ antenna with segments 7-13 with rhinaria and forming a club (Fig. 82), segment 3 longer 
than 4, 4-6 subequal in length, club segments more than 2 times as long as wide; cf antenna 15-segmented, 
filiform, segment 3 shorter than 4, 4 shorter than 5, 5-15 subequal in length. Head viewed frontally with 
eyes further apart than the height of an eye measured medially; inner orbits of face reticulate-rugose, the 
sculpture extending to and obscuring the malar grooves; occiput reticulate-rugose. Pronotal plate 
(Fig. 89); medial bridge broad; outer margins of pronotal plate angular; pronotum either side of pronotal 
plate pubescent. Scutellar fovea polished, shallow; scutellar cup sculptured, apex with a small fovea; 
scutellar disc with appearance of radiating sculpture, apex square. Mesopleuron finely colliculate above 
mesopleural suture, strigose below (Fig. 88); metapleuron weakly sculptured, polished, ridged on lower 
margins adjacent to front coxae; anteroventral cavity small, pubescent; propodeal carinae weakly bowed 
medially, densely pubescent on outer margins; nucha obscured by pubescence. Segment 2 of gaster the 
largest, occupying most of lateral area, apical half and whole of visible parts of segments 3 and 4 finely 
punctate; hypopygium broad apically, with sub-basal hairs or setae. Fore wing of female with a few hairs on 



AFROTROPICAL EUCOILIDAE 189 

veins, apical half bare, dotted with hair bases on under surface with a fringe of hairs on posterior angle, 
more pronounced on hind wing; radial cell of fore wing closed on wing margin though sometimes weakly 
pigmented; vein R { becoming thinner at junction with Rs 2 ; vein M distinct, Rs + M not apparent (Fig. 90); 
fore wing of male as in female except for apical fringes of hairs on margin. Coxae slightly longer than wide, 
swollen medially, weakly sculptured; trochanters longer than wide; femora long, weakly swollen medially, 
weakly sculptured; tibiae sculptured, pubescent; tarsi narrower than tibiae, weakly sculptured, metatarsus 
as long as remaining tarsal segments combined. Colour: antenna light brown; head and thorax black, 
except metapleuron, base of gaster and coxae chestnut brown; base and medial area of wings infuscated. 

Material examined 

Holotype $, Kenya: Nairobi, Karura, Furect, 13.xii.1970 (Stubbs) (BMNH). 

Paratypes. Kenya: 1 c? (BMNH). Zaire: 25 ?, 12 cf (MRAC). Zimbabwe: 2 $ (BMNH). South Africa: 
1 $ (BMNH). Uganda: 1 $> (BMNH). 

Eucoila veleda sp. n. 

Description. $ antenna with segments 6-13 with rhinaria and forming a club, segment 3 longer than 4, 4 
and 5 subequal in length, club segments 2 times as long as wide medially. Head viewed frontally with eyes 
further apart than the height of an eye measured medially; inner orbits of face with a trace of strigose 
sculpture, not extending to malar grooves; face with sparse scattered setae; occiput weakly strigose. 
Pronotal plate cf. Fig. 89; medial bridge narrow; outer parts of plate angulate (cf. Fig. 86); pronotum either 
side of pronotal plate pubescent. Scutellar fovea polished, deep; scutellar cup with a few transverse striae; 
scutellar disc cf. Fig. 87. Mesopleural suture with broken sculpture below (cf . Fig. 88); metapleuron ridged 
apically; anteroventral cavity large, pubescent; propodeal carinae bowed, densely pubescent on outer 
margins; nucha obscured by pubescence. Segment 2 of gaster the largest, occupying most of lateral area; 
tergites 3-4 partially visible ; apical third of 2 and visible parts of 3 and 4 finely punctate ; hypopygium , when 
visible, broad apically with sub-basal setae. Fore wing with a few scattered hairs on veins, apical half bare, 
dotted with hair bases, with a small fringe of hairs on posterior angle, prominent on hind wings; radial cell 
of fore wing open on wing margin; medial area of wing infuscate. Coxae slightly longer than wide, swollen 
medially, polished; trochanters a little longer than wide; femora long, weakly swollen basally, polished; 
tibiae narrow basally, widest apically with scattered setae; tarsi thinner than tibiae, metatarsus as long as 
remaining tarsal segments combined. Colour: antenna reddish brown; head and thorax black (except 
metapleuron reddish brown); gaster and legs orange-red. 
Cf unknown. 

Material examined 

Holotype g, Zimbabwe: Salisbury, i.1976 (Watsham) (BMNH). 

Paratypes. Zimbabwe: 1 $ (BMNH). Zaire: 3 $ (MRAC). Nigeria: 1 $ (BMNH). 

GLAURASPIDIA Thomson 

Glauraspidia Thomson, 1862: 307. Type species: Eucoela subtilis Dahlbom [= Cothonaspis microptera 

Hartig], by subsequent designation of Foerster, 1869: 351. 
Aglaotoma Foerster, 1869: 334. Type species: Cothonaspis codrina Hartig, by original designation. 

[Synonymised by Hellen, 1960: 9.] 
Apistophyza Foerster, 1869: 351. Type species: Cothonaspis microptera Hartig, by original designation. 

[Synonymised by Cameron, 1890: 24.] 
Diranchis Foerster, 1869: 360. Type species: Diranchis copulata Foerster, by original designation and 

monotypy. [Synonymised by Weld, 1952: 20.] 

Diagnosis. $ antenna 13-segmented, filiform-clavate, segments 9-13 usually with rhinaria, forming a 
weak club (Fig. 93); cf antenna filiform-flagelliform. Head viewed frontally longer than wide, frons viewed 
laterally protruding to form an angular shape, more so in males; occiput smooth and shining. Pronotal plate 
viewed frontodorsally with large fovea either side, open on lateral margins (Fig. 95), with dense woolly 
pubescence on either side. Mesopleuron smooth and polished, rarely finely coriaceous; mesopleural suture 
either absent or indistinct; scutellar cup raised above level of scutellar disc, disc reticulate-rugose to finely 
coriaceous, sometimes dull; propodeum viewed dorsally and laterally with dense woolly pubescence. Base 
of tergite 2 of gaster either with dense woolly pubescence or felt-like ring obscuring view of tergite 1 
(cf. Fig. 105). Wings pubescent with apical hair fringe; radial cell open in European species, closed in the 
Afrotropical ones. 

Distribution. Europe, Africa, Seychelles. 



190 J. QUINLAN 

Discussion. Nordlander (1982) treats Glauraspidia as belonging to his Chrestosema-group of genera, 
comprising Chrestosema, Glauraspidia, Pseudopsichacra, Odonteucoila, Dieucoila and Leptolamina. The 
relationships of the group are poorly understood because of the paucity of material. Nordlander recognizes 
three species of Glauraspidia in Europe. Two new species are here recorded from the Afrotropical region. 
For purposes of comparison the type species of the genus is also included in the key and is redescribed. 

Key to the type species and the Afrotropical species of Glauraspidia Thomson 

1 Radial cell of fore wing open on wing margin (Fig. 96) . 

Wing of female short, not extending past apex of gaster, of normal length in male; hair 
fringe short ; scutellar cup large , oval , disc reticulate-rugose microptera (Hartig) (p . 190) 

- Radial cell of fore wing closed on wing margin (Fig. 97) 2 

2 Face and occipital region smooth and shining; mesoscutum smooth and polished; scutellar cup 

elliptical, pitted, lateral margins almost imp unctate, apex reticulate scyphussp. n. (p. 191) 

- Inner orbits of face and occiput finely sculptured; mesoscutum polished, very finely coriaceous 

in upper medial and lateral regions; scutellar cup long and narrow, disc rounded apically, 
finely sculptured (Fig. 101) casca sp. n. (p. 190) 

Glauraspidia casca sp. n. 

(Figs 98-102) 

Description. $ antenna weakly subclavate, segment 3 subequal in length to 4+5, 4-8 each progressively 
shorter, 11-13 with pronounced rhinaria (Fig. 98); C? antenna 15-segmented, filiform, segments 3—13 
subequal in length, tapering apically (Fig. 99). Head viewed frontally with eyes further apart measured 
medially than the height of an eye; inner orbits with fine sculpturing extending from antennal sockets to 
about half way down eye margins; malar grooves obscured by broken striate sculpture; occiput finely 
sculptured, dense woolly pubescence on lateral margins. Pronotal plate with medial bridge narrow, foveae 
long and narrow (Fig. 100). Mesoscutum polished, finely coriaceous on upper medial and lateral surfaces, 
notauli absent; lateral bars of scutellum sculptured; scutellar foveae small, shallow; scutellar cup long and 
narrow, almost extending to apex of scutellar disc; disc rounded apically, surface almost smooth with fine 
broken sculpture (Fig. 101). Mesopleural suture absent; lower part of mesopleuron densely pubescent; 
anteroventral cavity obscure. Gaster with segment 2 the largest in lateral view, segments 3 and 4 partially 
visible in lateral view; apex of segment 2 and visible part of 3 coriaceous-punctate (Fig. 102); hypopygium 
pronounced. Wing surfaces densely pubescent, apical margins with a long hair fringe; radial cell of fore 
wing closed on wing margin; veins Rs + M and M weakly indicated. Colour: antenna yellow basally, apical 
segments darker; head, thorax and gaster chestnut brown; legs dark yellow. 

Material examined 
Holotype 5, Seychelles (David) (BMNH). 
Paratypes. Seychelles: 8 2,6c? (BMNH). 

Glauraspidia microptera (Hartig) 

(Figs 93-96) 

Cothonaspis microptera Hartig, 1840: 201. Holotype $, Germany (ZSBS) [examined]. 
Eucoela subtilis Dahlbom, 1842: 307. Holotype $, Sweden (ZI) [examined]. 

Description. 2 antenna with segments 1-8 filiform, 9-13 swollen medially with rhinaria, forming a club 
(Fig. 93), segment 3 very weakly curved, as long as 4+5, 5 and 6 subequal in length, 7 and 8 subequal, each 
shorter than 6; C? antenna 15-segmented, flagelliform, segment 3 curved, longer than 4+5 (Fig. 94), apical 
segments tapering. Head viewed laterally, particularly in female, wedge-shaped; eyes further apart 
measured medially than the height of an eye; malar grooves percurrent; face with scattered setae; occipital 
carina pronounced. Pronotal plate (Fig. 95). Mesoscutum smooth and polished, notauli absent, upper 
surface with scattered pubescence; lateral bars of scutellum weakly sculptured; scutellar foveae large, 
deep, smooth and polished; scutellar cup oval (but variable), almost extending to apex of scutellar disc, 
with a large median fovea; disc rounded apically, surface with reticulate-rugose-coriaceous sculpture, 
viewed dorsally appearing radiate in some specimens. Mesopleural suture absent. Carinae of propodeum 
sub-parallel, obscured by dense woolly pubescence on either side. Gaster with segment 1 obscured by 
dense woolly pubescence at base of tergite 2 (cf. Fig. 102); tergite 2 the largest in lateral view, remaining 



AFROTROPICAL EUCOILIDAE 191 

segments not visible ; ventral spine of gaster with sub-basal hairs. Wings with short apical hair fringe ; radial 
cell of fore wing open on wing margin; wings narrow. Colour: antenna orange-yellow; head, thorax and 
gaster blackish; legs orange-yellow. 

Distribution. Europe, not recorded from the Afrotropical region. 

Glauraspidia scyphus sp. n. 

(Figs 97, 103-105) 

Description. $ antenna moniliform, segment 3 as long as 4+5, 5-8 each shorter than 4, subequal to each 
other, apical 4 segments with rhinaria, forming a distinct club (Fig. 103); cf antenna 15-segmented, 
filiform, segment 3 weakly curved (Fig. 104), segment 5 swollen. Head viewed frontally with eyes slightly 
further apart measured medially than the height of an eye; cheeks converging; face with scattered hairs; 
mandibles tridentate; face smooth and polished; malar grooves percurrent; back of head smooth and 
polished. Mesoscutum smooth and polished, notauli absent, a row of scattered hairs in their place; lateral 
bars of scutellum weakly sculptured (cf. Fig. 101); scutellar cup elliptical; scutellar disc rounded apically, 
lateral surface almost impunctate, apex reticulate. Mesopleural suture absent; precoxal area of meso- 
pleuron densely pubescent; anteroventral cavity obscure, not hairy. Segment 1 of gaster short, wider than 
long, crenulate, partially obscured by the dense woolly pubescence on the propodeum and at base of tergite 
2; segment 2 the largest in lateral view, segments 3-5 visible in lateral view; apex of segment 2 and visible 
part of 3-5 punctate-coriaceous; hypopygium not pronounced (Fig. 105). Wing with apex broad; radial cell 
of fore wing closed on wing margin (Fig. 97); cubitus indicated, not pigmented. Colour: antenna yellow 
basally, apical segments darker; head brownish red, thorax and gaster light chestnut red; legs bright. 

Material examined 

Holotype $, Ivory Coast: Bouake, Bo'Pri, 10-13.iii.1984 (Matthews) (BMNH). 

Paratypes. Ivory Coast: 1 §, 1 cf (BMNH). Zaire: 10 $, 18 cf (MRAC). Zambia: 2 $ (BMNH). 
Zimbabwe: 2 $ (BMNH). 

Remarks. Distinguished from the European species microptera by the closed radial cell, the moniliform 
antenna of the female and the filiform antenna of the male. 

LEPTOPILINA Foerster 

Leptopilina Foerster, 1869: 348. Type species: Cothonaspis longipes Hartig, by original designation and 
monotypy. 

Diagnosis. $ antenna 13-segmented with variable number of club segments, each with rhinaria, generally 
clavate although sometimes slender and filiform; cf antenna 15-segmented, segment 3 shorter than 
following segments, flattened on outer margins (Fig. 106). Head narrow, eyes protruding; face viewed 
frontally smooth and polished with sparse hairs; malar grooves percurrent; mandibles tridentate; eyes 
further apart measured medially than height of an eye; viewed dorsally ocelli equally spaced, weakly 
raised. Pronotal plate weakly protruding, lateral foveae either side of pronotal plate broad, open on lateral 
margins. Mesoscutum smooth and polished, without trace of notauli, a line of hairs in their place; lateral 
bars of scutellum smooth and polished; scutellar fovea smooth and shallow (deep under lateral bars); 
scutellar disc variable in sculpture from punctate-reticulate to reticulate-rugose to nearly smooth with a few 
sparse rugae, sometimes more longitudinally directed (Fig. 107), apex rounded; scutellar cup (plate of 
Nordlander, 1982) variable in both size and shape, usually raised (cf. Figs 151, 159) smooth and shining to 
pitted with a large apical fovea. Mesopleuron smooth and polished; ventral border of pronotum 
sculptured; mesopleural suture distinct, straight or curved; metapleuron often with ridges extending from 
posterior margin; area below ventral margin of mesopleuron pubescent, anteroventral cavity bare; 
propodeal carinae distinct; propodeum not elongate, pubescence on either side dense, sculptured. 
Segment 1 of gaster short, wider than long, sculptured, crescent-shaped; base of segment 2 (petiole of 
Nordlander, 1982) widened posteriorly (Fig. 108), with a ring of hairs sometimes dense but broken on 
dorsal surface with a few long hairs on lateral margins behind ring of hairs (Fig. 108); tergite 2 the largest, 
occupying most of the area in lateral view; tergites 3-5 generally visible. Wings generally broad, rounded 
apically, surface pubescent, apical margins with fringe of hairs, radial cell of fore wing open or closed on 
wing margin. Legs of normal shape, variable between species; hind coxa with hairs on posterodorsal 
surface, sometimes tufted (cf. Fig. 109). Colour: antenna yellowish brown-black; head and thorax 
generally darker than gaster which is yellow to brown to blackish; legs yellowish. 



192 J. QUINLAN 

Discussion. Leptopilina received little attention (Weld, 1952) until Nordlander's (1980) major contribu- 
tion, which has considerably improved our understanding of the genus. Nordlander recorded Leptopilina 
from the Holarctic, Afrotropical and Oriental regions and gave a key to the five European species. In this 
study 14 species, 9 of them new, are recognised from the Afrotropical region. 

Key to the European and Afrotropical species of Leptopilina Foerster 

Females (the female of mahensis is unknown) 

1 Metapleuron without ridges in upper region; scutellar cup oval (Fig. 136), surface excavate or 

smooth, sometimes smooth with punctures 2 

- Metapleuron with or without ridges in upper region; scutellar cup elongate, surface convex to 

smooth , if oval , ridges present in upper region of metapleuron (Fig. 167) 4 

2 Mesoscutum with hairs present in place of the notauli; scutellar disc without a distinct rim 

(Fig. 136) c7av/pes(Hartig)(p. 195) 

- Mesoscutum smooth and polished without notauli or hairs in the same positions; scutellar disc 

with a distinct rim (Fig. 135) 3 

3 Antenna long and slender, segment 3 shorter than 4, club 8-segmented (Fig. 116), club 

segments 2 times as long as broad (Fig. 116). 

Pronotal plate with narrow medial bridge (cf . Fig. 159) fimbriata (Kieffer) (p. 196) 

- Antenna short, segment 3 longer than 4, club 7-segmented (Fig. 113), club segments broader 

than long longipes (Hartig) (p. 197) 

4 Antennal segment 3 clearly shorter than 4 (Figs 114, 117); metapleuron with ridges in upper 

region (Fig. 167) 5 

- Antennal segment 3 subequal to or longer than 4 (cf. Fig. 115); metapleuron with or without 

ridges in upper region 9 

5 Radial cell of fore wing open on wing margin (Fig. 141) ; antennal club 7-segmented. 

misensus sp. n. (p. 198) 

- Radial cell of fore wing closed on wing margin (cf. Fig. 150); antennal club 7-9-segmented, 

with rhinaria 6 

6 Antennal club 7-segmented (Figs 117, 118) 7 

- Antennal club 8-9-segmented (Figs 119, 120) 8 

7 Club segments of antenna at least 2 times as long as broad; scutellar cup broadest medially, 

tapering apically, not extending to apex of disc, with radiating reticulate-rugose sculpture 

(Fig. 169) thetussp. n.(p. 199) 

- Club segments of antenna at least 3 times as long as broad; scutellar cup elongate, almost oval, 

not extending to apex of disc, with fine reticulate-coriaceous sculpture (Fig. 162) ityssp. n. (p. 197) 

8 Antennal club 8-segmented (Fig. 119), apical segments of antenna dark; scutellar cup 

semi-oval, scutellar disc reticulate-rugose (Fig. 148); radial cell of fore wing closed on wing 
margin (Fig. 150) apella sp. n. (p. 194) 

- Antennal club 9-segmented (Fig. 120), apical segments of antenna pale; scutellar cup widest 

medially, converging apically, scutellar disc with radiating striae (Fig. 143) ; radial cell of fore 

wing open on wing margin fannius sp. n. (p. 195) 

9 Metapleuron without ridges in upper area (Fig. 121), a single ridge sometimes present 

posteriorly; antenna short, 0-5 times body length, segment 3 longer than 4, apical 4-5 
segments forming a club. 

Scutellar cup long and narrow; radial cell of fore wing closed, with a short radius. 

boulardi (Barbotin, Carton & Kelner-Pillault) (p. 195) 

- Metapleuron with 2 or more ridges in upper area (Fig. 157); antenna variable in proportions. ... 10 

10 Radial cell of fore wing open on wing margin (Fig. 122) 11 

- Radial cell of fore wing distinctly closed on wing margin (Fig. 140) 12 

11 Scutellar cup with pronounced rim, margins not converging apically (Fig. 123). 

atraticeps (Kieffer) (p. 194) 

- Scutellar cup without distinct rim, margins converging towards apex of disc. 

Radial cell (Fig. 158) faunus sp. n. (p. 196) 

12 Scutellar cup very broad medially, narrowing basally, converging towards apex of disc (e.g. 

Fig. 124) 13 

- Scutellar cup longer than wide , not converging towards apex of disc (Fig. 125) 15 

13 Metapleural ridges converging towards propodeum. 

Propodeal carinae parallel (cf. Fig. 149); radial cell (Fig. 140). 

heterotoma (Thomson) (p. 196) 



AFROTROPICAL EUCOILIDAE 193 

- Metapleural ridges parallel 14 

14 Scutellar cup viewed dorsally not extending to apex of disc; antenna, head and thorax pale 

yellow; radial cell of fore wing as broad as long (Fig. 129), open on wing margin. 

pisonissp. n. (p. 198) 

- Scutellar cup viewed dorsally extending to apex of disc (Fig. 127); antenna orange basally and 

apically, dark medially, thorax and gaster dark brownish orange; radial cell of fore wing 
elongate, 2 times as long as broad medially (Fig. 128), closed on wing margin. syphaxsp. n. (p. 199) 

15 Antennal club 7-segmented, sharply defined. 

Scutellar cup without conspicuous rim, almost oval, disc finely reticulate-rugose 
(Fig. 125); propodeal carinae widely separated, almost parallel; scutellar disc reticulate- 
rugose; metapleural carinae subparallel (Fig. 131); apical segments of gaster finely punctate. 

WcforiaeNordlander(p. 200) 

- Antennal club 8-segmented (filiform, club segments defined by presence of rhinaria) 

(Fig. 132) vestesp. n.(p. 200) 

Males (the males of atraticeps , itys and thetus are unknown) 

1 Metapleuron without ridges in upper region; scutellar cup oval, surface excavate or smooth, 

sometimes with punctures 2 

- Metapleuron with or without ridges in upper region; scutellar cup elongate, surface convex to 

smooth, if oval, ridges present in upper region of metapleuron (cf . Fig. 154) 4 

2 Mesoscutum smooth and polished without notauli or hairs in the same position; scutellar disc 

with a distinct rim (Fig. 135) 3 

- Mesoscutum with hairs in place of notauli (Fig. 110); scutellar disc without a rim (Fig. 110). 

clavipes (Hartig) (p. 195) 

3 Antennal segment 3 distinctly shorter than 4; cubitus distinctly pigmented. 

fimbriate (Kieffer) (p. 196) 

- Antennal segments 3 and 4 subequal in length (Fig. 113); cubitus absent. longipes (Hartig) (p. 197) 

4 Scutellar cup tapering sharply apically, broadest medially (Fig. 124) 5 

- Scutellar cup oval or if longer than broad not tapering apically, generally as broad apically as 

medially (Figs 125, 148) 8 

5 Radial cell of fore wing open or closed (Figs 112, 129); metapleural ridges not converging 

towards propodeum , almost parallel (Fig. 130) 6 

- Radial cell of fore wing closed (Fig. 140); metapleural ridges converging towards propodeum 

(Fig. 126) heterotoma (Thomson) (p. 196) 

6 Antennal segment 3 shorter than 4 (Fig. 144) 7 

- Antennal segments 3 and 4 subequal in length (Fig. 168). 

Scutellar cup extending to apex of scutellar disc (Fig. 127) syphax sp. n.(p. 199) 

7 Scutellar cup viewed dorsally not extending to apex of scutellar disc, disc reticulate-rugose 

(Fig. 143) ;pronotal plate (cf. Fig. 152) fannius sp. n.(p. 195) 

- Scutellar cup viewed dorsally not extending to apex of disc, with radiating striae ; pronotal plate 

(Fig. 166). 
Propodeal carinae converging; radial cell of fore wing open misensussp. n.(p. 198) 

8 Metapleuron without ridges in upper region; propodeal carinae closer together apically 

(Fig. 153); radial cell of fore wing closed on wing margin. 

boulardi (Barbotin, Carton & Kelner-Pillault) (p. 195) 

- Metapleuron with ridges in upper region; propodeal carinae variable; radial cell of fore wing 

open or closed on wing margin 9 

9 Scutellar cup oval (cf. Fig. 148); segment 3 of antenna shorter than 4. 

Veinifa -I- M of radial cell not indicated (Fig. 158) 10 

- Scutellar cup elongate ; segment 3 of antenna shorter than or subequal to 4 11 

10 Antennal segments 3 and 4 slender, curved (cf. Fig. 133); scutellar disc reticulate-rugose, 

scutellar cup large (cf . Fig. 123) faunus sp. n. (p. 196) 

- Antennal segments 3 and 4 robust and swollen (Fig. 133); scutellar disc with radiating strigose 

sculpture, scutellar cup small (cf . Fig. 148) mahensis (Kieffer) (p. 198) 

11 Antennal segments 3 and 4 subequal in length (Fig. 132) (4 weakly swollen) vesfasp. n.(p. 200) 

- Antennal segment 3 distinctly shorter than 4 (Fig. 147) 12 

12 Radial cell of fore wing with vein Rs + M in form of a distinctive spur (Fig. 146); antennal 

segment 4 swollen medially and curved, outer side flattened (Fig. 147). 

vicfor/aeNordlander(p. 200) 



194 J. QUINLAN 

- Radial cell of fore wing without a distinctive spur (Fig. 150) ; antennal segment 4 weakly swollen 

medially and curved, outer side flattened. 

Pronotal plate (Fig. 159) 13 

13 Scutellar disc reticulate-rugose (Fig. 148), rim of scutellar cup thick, not converging apically; 

metapleural ridges parallel apella sp. n. (p. 194) 

- Scutellar disc coarsely rugose-reticulate, scutellar cup converging apically (cf. Fig. 124); 

metapleural ridges weakly converging towards propodeum pisonissp. n. (p. 198) 

Leptopilina apella sp. n. 

(Figs 119, 148-150, 159) 

Description. $ antenna weakly clavate, segments 6-13 with rhinaria, segment 3 shorter than 4 (Fig. 119); 
Cf antenna with segments 3-4 curved (cf. Fig. 147). Eyes large, converging towards clypeal area, closer 
together than the height of an eye where they converge; malar grooves fine. Pronotal plate with medial 
bridge narrow, as wide as a fovea (Fig. 159). Lateral bars of scutellum broad-based, triangular; scutellar 
fovea smooth and shining, shallow, lenticular; scutellar cup small, oval, not extending to apex of disc; 
scutellar disc reticulate-rugose, apex rounded (Fig. 148). Mesopleural suture converging towards meta- 
pleuron with two ridges in upper region (cf. Fig. 167); anteroventral cavity pubescent; lateral margins of 
propodeum densely pubescent on under margins of carinae, weakly sculptured; propodeal carinae parallel 
(Fig. 149). Segment 1 of gaster in form of short crenulate ring; segment 2 with tuft of pubescence basally on 
lateral margins; segment 3 partially visible; gaster impunctate. Radial cell of fore wing closed on wing 
margin, broad apically; vein 3r-ra produced; vein M indicated basally, weakly pigmented (Fig. 150). Legs 
long and slender; hind coxa with tuft of hairs on posterodorsal margin. Colour: antenna yellow basally, 
segments 6-13 blackish; head blackish; thorax chestnut brown; gaster brownish yellow; legs yellow. 

Material examined 
Holotype J, Madagascar: Tarn, Perinet, 27.iv.-3.v.l983 (Noyes & Day) (BMNH). 
Paratypes. Madagascar: 1 2, 3 cf (BMNH). Zaire: 8 $ (MRAC). 

Leptopilina atraticeps (Kieffer) 

(Figs 122, 123, 151) 

Ectolyta atraticeps Kieffer, 1911: 312. Holotype $, Seychelles (BMNH) [examined]. 

Description. $ antenna clavate, segments 7-13 forming a club, rhinaria weakly indicated, each segment 
more than 2 times as long as broad, segment 3 slightly longer than 4, less than 4+5,4 longer than 5. Eyes 
oval, as far apart measured medially as the height of an eye; subantennal sutures absent. Pronotal plate 
projected forward; anterior part finely sculptured; lateral foveae open on either side of medial bridge, wide 
(Fig. 151). Mesoscutum smooth and polished, variable surface sculpture indicated through the chitin 
(Fig. 123); lateral bars of scutellum smooth and polished; scutellar foveae deep, polished; scutellar cup 
elliptical, widest medially, with a wide rim, with a large apical fovea; scutellar disc reticulate-rugose, 
rounded apically. Mesopleuron smooth and polished; mesopleural suture distinct, curved; metapleuron 
smooth and polished with 2 conspicuous ridges; anteroventral cavity obscured; sides of propodeum with 
dense short pubescence; propodeum almost parallel, very weakly bowed medially. Segment 1 of gaster in 
form of crenulate ring, wider than long; segment 2 with a dense ring of short hairs basally not completed on 
dorsal surface; tergites 3 and 4 partially visible. Wings not conspicuously narrow; pubescent with long 
apical hair fringe; radial cell of fore wing partially open on wing margin; veins Rs + M and M weakly 
indicated (Fig. 122). Legs slender; hind coxa with a tuft of pubescence postero-dorsally. Colour: antenna 
yellowish orange, some median segments light brown; head reddish brown; thorax and gaster orange- 
yellow; legs orange-yellow. 
Cf unknown. 

Material examined 
Holotype $, Seychelles: Mahe I., Mare au Cochons (BMNH). 
Paratype. Nigeria: 1 § (BMNH). 

Remarks. Similar in many respects to faunus, but differs in the shape of the radial cell and the scutellar cup. 



AFROTROPICAL EUCOILIDAE 195 

Leptopilina boulardi (Barbotin, Carton & Kelner-Pillault) 
(Figs 115, 121, 152, 153) 

Charips mahensis Kieffer, 1911: 313. Holotype cf , Seychelles (BMNH) [examined]. [Junior secondary 
homonym in Leptopilina of Erisphagia mahensis Kieffer (see p. 198).] [Synonymised by Nordlander, 
1980: 432.] 

Cothonaspis (Cothonaspis) Boulardi Barbotin, Carton & Kelner-Pillault, 1979: 20. Holotype 9, Guade- 
loupe (MNHN) [not examined]. 

Description. $ antenna clavate, the 4 or 5 apical segments forming a club, each of club segments with 
rhinaria, each a little longer than broad, generally darker than the yellow basal segments, segment 3 as long 
as 4+5, 4 longer than 5, 5 and 6 subequal in length, 7 longer than 6 (Fig. 115); d" antenna 15-segmented, 
filiform, segment 3 shorter than 4, 4 curved, swollen, longer than 5. Eyes almost round, further apart than 
the height of an eye measured medially; subantennal sutures prominent on lower face, sparse hairs present; 
malar grooves percurrent. Pronotal plate projected forward; lateral foveae long and narrow (Fig. 152). 
Mesoscutum smooth and polished, notauli absent; lateral bars of scutellum smooth and polished; scutellar 
foveae shallow; scutellar cup long and narrow, widest medially, apex weakly converging, apex with a large 
fovea, area in front weakly convex, sculptured; scutellar disc with radiating ridges, apex of disc rounded. 
Mesopleuron smooth and polished; mesopleural suture distinct, curved; metapleuron with one or two 
incomplete ridges (Fig. 121); nucha short, weakly pubescent; propodeal carinae closer together in upper 
half, bowed medially. Segment 1 of gaster crenulate, in form of a crescent; segment 2 widened dorsally, 
hairy ring at base very thin, only a few hairs present laterally; visible parts of segments 3-5 punctate. Fore 
wings not conspicuously narrow, pubescent, apical hair fringe long; radial cell of fore wing closed on 
margin; 2r-m distinct; Rs + M and M weakly indicated. Legs robust; hind coxa without a tuft of hairs 
postero-dorsally. Colour: apical segments of antenna brownish black, basal segments yellow; head, thorax 
and gaster brownish black-chestnut; legs yellow. 

Material examined 

Charips mahensis Kieffer, holotype cf , Seychelles (Percy Sladen Trust Expedition) (BMNH). 

Aldabra: 1 $ (BMNH). Guadeloupe: 1 $, 1 cf (BMNH). Madagascar: 4 ? (BMNH). South Africa: 10 $ 
(BMNH). Zaire: 32 $, 33 cf (MRAC). Zambia: 4 $ (BMNH). Zimbabwe: 2 $, 1 cf (BMNH). 

Remarks. Separated from all other species in the genus by the absence of metapleural ridges, form of 
antenna and scutellar cup. It is a solitary primary internal parasite of Drosophila melanogaster Meigen (see 
Nordlander, 1980). 

Leptopilina clavipes (Hartig) 

(Figs 110, 136) 

Cothonaspis clavipes Hartig, 1841: 357. 

Remarks. This species is included in the key for comparative purposes. Nordlander (1980) gives a 
description and synonymy. At present clavipes is known to occur in Europe and North America but it has 
not been found in the Afrotropical region. 

Leptopilina fannius sp. n. 

(Figs 120, 143, 144, 154, 155, 156) 

Description. 2 antenna filiform, segments 5-13 with rhinaria, segment 3 shorter than 4, subequal to 5, 
apical two segments pale (Fig. 120); cf antenna 15-segmented, filiform, segment 3 shorter than 4, weakly 
curved (Fig. 144). Eyes oval, as far apart measured medially as the height of an eye; face smooth and 
polished; malar grooves percurrent. Pronotal plate weakly protruding; medial bridge broad (cf. Fig. 159). 
Mesoscutum smooth and polished; lateral bars of scutellum broad basally, short; scutellar cup longer than 
wide, not extending to apex of disc, widest medially, converging apically (Fig. 143); scutellar disc 
reticulate-rugose laterally, rounded apically (sometimes the margins are striated). Mesopleuron smooth 
and polished; mesopleural suture almost straight; metapleuron with three distinct ridges, the upper ridge 
branched (Fig. 154); anteroventral cavity large, hairless; lateral margins of propodeum crenulate, sparsely 
pubescent; propodeal carinae bowed (cf. Fig. 161). Segment 1 of gaster in form of crenulate ring, wider 
than long; tergite 2 with a few sparse hairs on lateral margins of base, apical half densely punctate; visible 
parts of segments 3 and 4 punctate. Fore wing relatively broad, rounded apically, surfaces pubescent, 
apical margins with a fringe of hairs; radial cell of fore wing closed on margin, weakly pigmented, broad 



196 J. QUINLAN 

apically ; vein 2r-m not projecting, merging into vein M; vein Rs + M not indicated (Fig. 155). Legs robust; 
hind coxa stout, a few hairs present on postero-dorsal margin. Colour: antenna yellow basally, dark 
medially, apical two segments pale yellow; head and thorax black, except metapleuron orange-brown; legs 
orange-yellow. 

Material examined 

Holotype $ , Zaire: Massif Ruwenzori, Kyandolire, 1700m, Camp des Gardes, 7- 15.x. 1952 (Vanschuyt- 
broeck & Kekenbosch) (MRAC) . 

Paratypes. Zaire: 1 $ , 6 cT (MRAC). 

Remarks. Separated from vesta by the relative proportions of the antennal segments and the form of the 
scutellar cup. 

Leptopilina faunus sp. n. 

(Figs 157, 158) 

Description. 2 antenna clavate, segments 7-13 with rhinaria, segment 3 longer than 4, 5 and 6 subequal, 
each shorter than 4 (cf. Fig. 156); C? antenna 15-segmented, segment 3 shorter than 4, both slender and 
curved (cf . Fig. 137). Eyes almost round, further apart measured medially than the height of an eye; malar 
grooves percurrent; face smooth and polished, a few sparse hairs present in clypeal region. Pronotal plate 
weakly protruding; lateral foveae open (but not readily appreciated); medial bridge broad (cf. Fig. 151). 
Mesoscutum smooth and polished; lateral bars of scutellum broad basally, longer than wide; scutellar 
foveae lenticular, smooth and shallow; scutellar cup elongated, not extending to apex of disc, widest 
medially, tapering apically, apical fovea large; scutellar disc reticulate-rugose, rounded apically. Meso- 
pleuron smooth and polished; mesopleural suture weakly curved; metapleuron with two strong ridges in 
upper quarter converging towards margins of propodeum (Fig. 157); anteroventral cavity pubescent; 
lateral margins of propodeum sculptured, partially covered with pubescence; propodeal carinae widely 
spaced, weakly bowed. Segment 1 of gaster in form of crenulate ring; tergite 2 with a few hairs at lateral 
margins of base, apical margins sparsely punctate; segments 3 and 4 partially visible, punctate. Fore wing 
broad, wing surfaces pubescent with apical hair fringe; radial cell of fore wing partially open on wing 
margin; vein 2r-m weakly projecting; veins M and Rs + M weakly indicated by pigmentation (Fig. 158). 
Legs slender; hind coxa with a few hairs on posterodorsal margin. Colour: antenna orange-yellow basally, 
brownish yellow apically, apical segment lighter than rest; head and thorax brown, metapleuron orange- 
brown; gaster chestnut brown; legs orange-yellow. 

Material examined 

Holotype $, Zaire: Nyasheke (Vole. Nyamuragira) , 1820 m, 14-26.vi.1935 (de Witte) (MRAC). 
Paratypes. Zaire: 3 5, 5 cf (MRAC). 

Remarks. Distinguished from atraticeps by the shape of the scutellar cup and radial cell. 

Leptopilina Hmbriata (Kieffer) 

(Figs 111, 116, 135, 137, 138) 

Eucoela fimbriata Kieffer, 1901: 175. 

Remarks. This species is included in the key for comparative purposes. Nordlander (1980) gives a full 
synonymy, description and bionomic information. It has not been found outside Europe. 

Leptopilina heterotoma (Thomson) 
(Figs 124, 126, 140, 160) 

Eucoila heterotoma Thomson, 1862: 403. Holotype $, Sweden (ZI) [examined]. 

Ganaspis monilicornis Kieffer, 1904: 622. Lectotype 2, France (MA) [not examined]. [Synonymised by 
Nordlander, 1980: 430.] 

Ganaspis subnuda Kieffer, 1904: 64. Holotype cf , Canary Is (not located). [Synonymised by Nordlander, 
1980: 430.] 

Erisphagia philippinensis Kieffer, 1916: 282. Lectotype $ , Los Banos (MNHN) [not examined]. [Synony- 
mised by Nordlander, 1980: 430.] 

Pseudeucoila bochei Weld, 1944: 65. Holotype $, U.S.A. (USNM) [examined]. [Synonymised by 
Nordlander, 1980:461.] 



AFROTROPICAL EUCOILIDAE 197 

Description. $ antenna weakly clavate, segments 3 and 4 subequal in length, 5 shorter than 4, apical 7 
segments with rhinaria forming a club (Fig. 160); cf antenna 15-segmented, filiform, segment 3 shorter 
than 4, 4 twisted and swollen medially (Fig. 137). Face smooth and polished with sparse hairs; anterior 
tentorial pits minute; malar grooves finely indicated; head viewed dorsally as wide as thorax; eyes large; 
occipital carinae distinct. Pronotal plate projected forward; medial bridge broad; anterior plate sculp- 
tured ; pronotum either side of plate pubescent. Mesoscutum smooth and polished without trace of notauli ; 
lateral bars of scutellum smooth and polished; scutellar fovea deep, lenticular; scutellar disc smooth to 
reticulate-rugose with scattered hairs, rounded apically ; scutellar cup smooth and polished, convex in front 
of apical pit, long, widest above medial area, converging apically and narrow (Fig. 124). Mesopleuron 
smooth and polished; mesopleural suture distinct, weakly curved; metapleuron depressed above ridges 1 
and 3, ridges 1, 2 and 3 entire (Fig. 126); lateral margins of propodeum below metanotal plate pubescent; 
propodeal carinae sub-parallel, wide (cf. Fig. 150). Segment 1 of gaster in form of a narrow ring, 
sculptured, weakly inclined posteriorly; tergite 2 with ring of dense pubescence ventrally, not complete on 
dorsal surface; tergite 3 partially visible; apical margins of tergite 2 and whole of visible parts of 3 punctate 
(cf . Fig. 108) . Wing surfaces pur oscent, apical margins with fringe of hairs; radial cell of fore wing closed on 
margin; vein Rs as long as vein 2r; vein M indicated, not pigmented (Fig. 140). Legs slender; hind coxa with 
distinct tuft of hairs on posterodorsal surface. Colour: antenna yellow basally, darker apically; head black; 
thorax and gaster dark brown-black. 

Material examined 
Eucoila heterotoma Thomson, holotype $ , Sweden: Skane, Baggeboda (ZI). 
Madagascar: 1 $ (MRAC). Palestine: 1 ?, 1 cT (BMNH). Zaire: 1 $ (MRAC). 

Remarks. The specimens from Madagascar and Palestine are only tentatively placed as this species 
because the apical segments of the antenna are white. L. heterotoma is world-wide in distribution 
(Nordlander, 1980). The species is a solitary primary internal parasitoid of Drosophila larvae and it has 
been widely studied in Europe and the U.S.A. 

Leptopilina ityssp. n. 

(Figs 118, 161,162) 

Description. $ antenna weakly clavate, segments 7-13 with rhinaria, 4 times as long as broad, segment 3 
shorter than 4, 4-6 subequal in length (Fig. 118). Eyes large, almost round, further apart measured 
medially than the height of an eye; face smooth and polished with sparse scattered hairs; malar grooves 
very fine, percurrent. Pronotal plate protruding; medial bridge as wide as a lateral fovea. Mesoscutum 
smooth and polished; lateral bars of scutellum broad-based, triangular; scutellar fovea wider than long, 
shallow; scutellar cup elongate, not extending to apex of disc, with a large apical fovea; scutellar disc finely 
reticulate-rugose, apex rounded (Fig. 162). Mesopleuron smooth and polished; mesopleural suture almost 
straight; metapleuron with two ridges in upper region, parallel, widely spaced (cf. Fig. 131); anteroventral 
cavity large, pubescent; lateral margins of propodeum sculptured, partially concealed by pubescence; 
propodeal carinae closer together anteriorly (Fig. 161). Segment 1 of gaster in form of a crescent-shaped 
crenulate ring, partially visible; lateral margins of base of tergite 2 with a ring of hairs, thin dorsally, thicker 
ventrally; tergites 3-5 partially visible in lateral view, sparsely punctate. Fore wing relatively narrow 
compared with other species in the genus, wing surfaces pubescent with apical hair fringe ; radial cell of fore 
wing closed on wing margin; vein 2r-m weakly protruding; veins M and Rs + M not indicated. Legs long 
and slender; hind coxa with a few short hairs on posterior dorsal margin. Colour: antenna yellow basally, 
apical 5-6 darker; head, thorax and gaster dark brownish black; legs yellow. 

Cf unknown. 
Material examined 

Holotype $>, Zimbabwe: Salisbury, Chishawasha, hi. 1980 (Watsham) (BMNH). 

Paratype. Zaire: 1 £ (MRAC). 

Remarks. The length of the antennal segments, shape of the scutellar cup and the sculpture of the scutellar 
disc separate this species from thetus. 

Leptopilina longipes (Hartig) 
(Figs 106-109, 113, 139) 

Cothonaspis longipes Hartig, 1841: 356. 

Remarks. This European species is included in the key for comparative purposes. It is the type species of 
the genus. Nordlander (1980) gives a description. 



198 J. QUINLAN 

Leptopilina mahensis (Kieffer) 

(Figs 133, 163, 164) 

Erisphagia mahensis Kieffer, 1911: 312. Holotype d\ Seychelles (BMNH) [examined]. 

Description. $ unknown. 

Cf antenna 15-segmented, filiform, segment 3 shorter than 4, weakly swollen apically, 4 curved and 
swollen medially (Fig. 133). Face smooth and polished; eyes large, protruding; anterior tentorial pits 
distinct; malar grooves percurrent but finely indicated; a few sparse hairs present; head viewed dorsally as 
wide as thorax; occipital carinae pronounced. Pronotal plate projected forward: posterior plate angled; 
medial bridge narrow; lateral foveae partially open (cf. Fig. 149). Mesoscutum smooth and polished, 
without trace of notauli or hairs in their place; lateral bars of scutellum polished; scutellar foveae large, 
deep and smooth; scutellar disc rounded apicallv, surface reticulate-rugose; scutellar cup almost round 
apically, narrower basally with a pronounced r j\, apical third with a large pit or fovea (cf. Fig. 148). 
Mesopleuron smooth and polished; mesopleural suture percurrent, almost straight; metapleuron polished 
and strongly ridged in anterodorsal corner; antero ventral cavity sculptured; propodeum not elongated in 
lateral view; nucha distinct; viewed dorsally propodeal carinae almost parallel, weakly bowed medially 
(Fig .163). Segment 1 of gaster in form of ring or collar , crenulate , widened posteriorly ; tergite 2 occupying 
the whole area of gaster in lateral view with a few hairs basally on lateral margins. Wing surfaces pubescent 
with apical hair fringe long; radial cell of fore wing closed, longer than wide; veins Rs + M and M absent 
(Fig. 164). Legs of normal shape and proportions; coxae with sparse hairs, hind coxa without a tuft of hairs. 
Colour: antenna orange-yellow; head and thorax light brown; gaster chestnut red; legs yellow. 

Material examined 
Holotype cf, Seychelles: Mahe I., Mare Cochons district, 1000-2000 ft, 26.i.-2.ii.l909 (BMNH). 

Remarks. Because the female is unknown the affinities of this species are uncertain. The male is separated 
from faunus by antennal, scutellar disc and radial cell characters. 

Leptopilina misensus sp. n. 

(Figs 114, 141,165-167) 

Description. $ antenna filiform, segments 5-13 with weak rhinaria, segment 3 shorter than 4, 4 subequal 
to 5, 6-13 subequal, apical segments lighter than rest of antenna (Fig. 114); cf antenna 15-segmented, 
filiform, segment 3 shorter than 4, 4 weakly swollen medially, longer than 5 (Fig. 165). Eyes almost round, 
further apart than the height of an eye measured medially; face smooth and polished; malar grooves 
percurrent. Pronotal plate weakly projecting; lateral foveae long and narrow (Fig. 166). Mesoscutum 
smooth and polished, without trace of notauli; lateral bars of scutellum smooth and polished; scutellar 
foveae broad basally; scutellar cup longer than wide with apical fovea widest medially, tapering or 
converging apically; scutellar disc with aberrant reticulate-rugose sculpture, rounded apically. Meso- 
pleuron smooth and polished; mesopleural suture distinct, weakly curved; metapleuron with 2 ridges, 
almost parallel; anteroventral cavity large, hairless (Fig. 167); lateral margins of propodeum crenulate; 
propodeal carinae parallel (cf. Fig. 163). Segment 1 of gaster in form of crescent-shaped crenulate ring; 
tergite 2 occupying almost the whole visible area in lateral view, with a few hairs on the lateral margins 
basally; tergites 3 and 4 partially visible; apex of tergite 2 and visible parts of 3 and 4 punctate. Fore wing 
relatively broad, apical margins rounded, with a fringe of hairs, wing surfaces pubescent; radial cell of fore 
wing open on wing margin, broad apically; vein 2r-m weakly projecting; Rs + M not indicated (Fig. 141). 
Legs slender; hind coxa without a tuft of hairs posterodorsally. Colour: antenna dark yellow basally, 
brownish medially, apical segments sometimes lighter; head, thorax and gaster brownish yellow. 

Material examined 
Holotype $, Zaire: Nyasheke (vole. Nyamuragira), 1820m, 14-26. [?month]. 1935 (de Witte) (MRAC). 
Paratypes. Zaire: 3 $, 18 cT (MRAC). Uganda: 2 C? (BMNH). 

Remarks. This species has characters in common with heterotoma in the scutellar cup, but the antennal 
characters and open radial cell enable it to be recognised easily. 

Leptopilina pisonis sp. n. 

(Figs 129, 130) 

Description. $ antenna clavate, segments 7-13 with rhinaria, swollen medially, segment 3 longer than 4, 4 
longer than 5, 5 and 6 subequal (cf. Fig. 156); C? antenna filiform, segment 3 shorter than 4, 4 swollen 



AFROTROPICAL EUCOILIDAE 199 

medially (cf. 137). Eyes oval, further apart measured medially than the height of an eye; face smooth and 
polished; malar grooves percurrent. Pronotal plate weakly protruding; medial bridge as wide as a lateral 
fovea (cf. Fig. 159). Mesoscutum smooth and polished, without trace of notauli; lateral bars broad, short 
and triangular; scutellar foveae lenticular, shallow; scutellar cup elongate, broadest medially, weakly 
converging apically, not extending to apex of disc; scutellar disc with large reticulate-rugose sculpturing, 
apex rounded (cf. Fig. 124). Mesopleuron smooth and polished; mesopleural suture weakly curved; 
metapleuron weakly ridged in upper quarter (Fig. 130); anteroventral cavity pubescent; lateral margins of 
propodeum partially obscured by dense pubescence, visible part crenulate; propodeal carinae weakly 
bowed (cf . Fig. 163) . Segment 1 of gaster in form of crescent-shaped crenulate ring; lateral margins of base 
of tergite 2 with dense pubescence ventrally, thin dorsally; tergites 3-6 partially visible in lateral view; 
apical margin of tergite 2 and whole of visible parts of 3-6 punctate. Fore wing moderately broad, wing 
surfaces pubescent with apical hair fringe; radial cell of fore wing partially open on wing margin; vein 2r-m 
not projecting; veins M and Rs + M not indicated (Fig. 129). Legs long and slender; hind coxa with a tuft of 
hairs on posterodorsal margin. Colour: antenna pale yellow basally , darkish medially, pale yellow apically; 
head, thorax and gaster orange-brown; legs pale yellow. 

Material examined 

Holotype $, Zaire: Eala, v. 1935 {Ghesquiere) (MRAC). 
Paratypes. Zaire: 30 g, 8 cf (MRAC). 

Remarks. Similar to syphax but differing in the shape and size of the scutellar cup, the radial cell and the 
colour pattern. 

Leptopilina syphax sp. n. 

(Figs 127, 128, 134, 142, 168) 

Description. $ antenna clavate, segments 7-13 subequal in length with rhinaria, 3 and 4 subequal in 
length, 5 shorter than 4, 6 shorter than 5, apical three segments lighter than median segments (Fig. 168); cf 
antenna 15-segmented, filiform, 3-5 subequal in length (Fig. 142). Eyes oval, as far apart as the height of 
an eye measured medially; face smooth and polished; malar grooves percurrent. Pronotal plate projecting; 
medial bridge broad. Mesoscutum smooth and polished, without trace of notauli; lateral bars of scutellum 
long, broad basally; scutellar cup longer than wide, extending to apex of disc, widest medially, converging 
apically (Fig. 127), apical fovea large; scutellar disc areolate-reticulate-rugose, rounded apically (Fig. 127). 
Mesopleuron smooth and polished; mesopleural suture almost straight; metapleuron with two ridges, 
weakly converging towards the propodeum; anteroventral cavity large, hairless (Fig. 134); lateral margins 
of propodeum canaliculate, weakly pubescent; propodeal carinae obscured by pubescence. Segment 1 of 
gaster in form of a crenulate ring, wider than long; tergite 2 with a few hairs on lateral margins basally; 
tergites 3-5 partially visible ; apex of tergite 2 and visible parts of 3-5 punctate . Fore wing relatively broad , 
rounded apically, wing surfaces pubescent with apical hair fringe; radial cell of fore wing closed on wing 
margin; vein 2r-m weakly projecting; vein M complete but faintly represented; vein Rs + M not indicated. 
Legs slender; hind coxa without a tuft of hairs posterodorsally. Colour: basal segments of antenna 
orange-yellow, medial segments brownish, apical three segments light yellow; head and thorax blackish; 
metapleuron and gaster orange-brown; legs orange-yellow. 

Material examined 
Holotype $, Zaire: R. Kilalamatambo (affl. Lusinga), 17.vii.1945 (de Witte) (MRAC). 
Paratypes. Zaire: 12 $, 24 cf (MRAC). 

Remarks. This species is closely related to pisonis but characteristics in the radial cell, scutellar cup and 
colour pattern enable them to be separated. 

Leptopilina thetussp. n. 
(Figs 112, 117, 169) 

Description. $ antenna weakly clavate, segment 3 shorter than 4, longer than 5, 5 and 6 subequal in 
length, 7-13 three times as long as broad, with rhinaria (Fig. 117). Eyes large, round, as far apart as the 
height of an eye measured medially; face smooth and polished; malar groove not visible; a few hairs present 
in clypeal area. Pronotal plate weakly projecting; medial bridge narrow, not as wide as a lateral fovea (cf. 
Fig. 159). Mesoscutum smooth and polished; lateral bars of scutellum broad basally, wide apically; 
scutellar foveae lenticular, smooth and shallow; scutellar cup extending almost to apex of scutellar disc, 
longer than wide, widest medially, converging apically (Fig. 169), apical fovea large, rim of scutellar cup 



200 J. QUINLAN 

thick; scutellar disc with radiating reticulate-rugose sculpture, apex rounded. Mesopleuron smooth and 
polished; mesopleural suture weakly curved; metapleuron with two widely spaced parallel ridges in upper 
region; anteroventral cavity large, a few setae present; lateral margins of propodeum obscured by dense 
pubescence; propodeal carinae parallel. Segment 1 of gaster in form of crenulate ring, wider than long; 
tergite 2 with a tuft of hairs on lateral margins, thinner on dorsal surface; tergites 3-5 partially visible; apex 
of segment 2 and visible parts of 3-5 with sparse punctures. Fore wing moderately broad, wing surfaces 
pubescent with apical hair fringe; radial cell of fore wing closed on wing margin; vein 2r-m weakly 
projecting; veins M and Rs + M not indicated (Fig. 112). Legs short; hind coxa with a few hairs on 
posterodorsal surface. Colour: antenna pale yellow; head, thorax and gaster pale chestnut brown; legs 
orange-yellow. 
Cf unknown. 

Material examined 

Holotype $ , South Africa: Port St John, Pondoland, 10-31.vii.1923 (Turner) (BMNH). 
Paratypes. South Africa: 2 2 (BMNH). Zaire: 3 5 (MRAC). 

Remarks. This species is separated from closely related species by the form of the antenna and the 
elongated scutellar cup. 

Leptopilina vesta sp. n. 

(Figs 132, 145, 170) 

Description. $ antenna filiform, segments 3-13 with rhinaria, segment 3 shorter than 4, 4 and 5 subequal 
in length (Fig. 132); cf antenna 15-segmented, filiform, segments 3 and 4 subequal in length, 4 weakly 
swollen (Fig. 145). Eyes large, round, as far apart measured medially as height of an eye; face smooth and 
polished; malar grooves finely indicated. Pronotal plate protruding; medial bridge as broad as a lateral 
fovea (cf. Fig. 159). Mesoscutum smooth and polished; lateral bars of scutellum triangular, short; scutellar 
foveae lenticular; scutellar cup longer than wide, not extending to apex of disc; scutellar disc reticulate- 
rugose, apex rounded (cf. Fig. 169). Mesopleuron smooth and polished; mesopleural suture converging 
towards metapleuron; metapleuron with three distinct ridges in upper region (cf. Fig. 134); anteroventral 
cavity pubescent (cf. Fig. 134); lateral margins of propodeum partially obscured by dense pubescence; 
propodeal carinae bowed (cf. Fig. 163). Segment 1 of gaster in form of narrow ring, weakly crenulate; 
tergite 2 with a tuft of pubescence basally on lateral margins; apex of tergite 2 and visible part of tergite 3 
finely punctate. Fore wing broad, apex rounded, wing surfaces pubescent with apical fringe of hairs; radial 
cell of fore wing closed on wing margin , narrow apically ; vein 2r-m protruding ; veins M and Rs + M weakly 
indicated, not pigmented (Fig. 170). Legs long and slender; hind coxa without tuft of hairs on posterodorsal 
margin. Colour: antenna yellowish basally, merging to brownish yellow apically; head and thorax dorsally 
blackish; mesopleuron and gaster brownish orange; legs yellow. 

Material examined 
Holotype $>, Cameroun: Nkoemvon, 1980 (Jackson) (BMNH). 
Paratypes. Cameroun: 1 cf (BMNH). Zaire: 2 $, 8 cf (MRAC). 

Remarks. The filiform antenna separates this species from all others in this genus. 

Leptopilina victoriae Nordlander 

(Figs 125, 131, 146, 147) 

Leptopilina victoriae Nordlander, 1980: 447. Holotype $, Seychelles (Nordlander coll.) [not examined]. 

Description. $ antenna clavate, segments 7-13 with rhinaria forming a club, segment 3 longer than 4, 5 
and 6 subequal, each shorter than 4 (cf. Fig. 156); cf antenna 15-segmented, segment 3 shorter than 4, 4 
swollen medially, bent, outer surface flattened (Fig. 147). Eyes large, almost round, further apart 
measured medially than the height of an eye; face smooth and polished with sparse hairs around clypeal 
area; malar grooves fine, percurrent. Pronotal plate protruding; lateral foveae round, narrow; medial 
bridge slightly broader than a fovea. Mesoscutum smooth and polished; lateral bars of scutellum short, 
broad based; scutellar cup elongate, weakly converging apically with a large apical fovea; scutellar disc 
rounded apically, surface reticulate-rugose (Fig. 125). Mesopleural suture directed strongly down towards 
margin of anteroventral cavity; metapleuron with three ridges (Fig. 131); anteroventral cavity hairless; 
lateral margins of propodeum crenulate, partially obscured by dense pubescence; propodeal carinae very 
weakly bowed, almost parallel (cf. Fig. 149). Segment 1 of gaster in form of a crescent, crenulate; lateral 



AFROTROPICAL EUCOILIDAE 201 

margins of base of tergite 2 with a thin ring of hairs and a few longer hairs behind them; tergites 3-5 
partially visible, impunctate. Fore wing moderately broad apically, wing surfaces pubescent, apical 
margins with a hair fringe; radial cell of fore wing closed on wing margin, broad apically; vein 2r-m 
projected; vein Rs + M not indicated; vein M weakly indicated by pigmentation (Fig. 146). Legs slender; 
hind coxa swollen, with a tuft of hairs on posterodorsal margin. Colour: antenna yellow basally, apical 
segments darker; head, thorax and gaster dark chestnut brown; legs orange-yellow. 

Material examined 

Seychelles: 14 $, 46 cf (including 1 $, 1 O" paratypes of L. victoriae Nordlander, labelled ES 551 and 
552) (BMNH). Uganda: 2 $ (BMNH). Zaire: 32 $, 18 cf (MRAC). South Africa: 1 2 (BMNH). 

Remarks. Separated from vesta by antennal, pronotal plate and scutellar cup characters. Nordlander 
(1980) considers victoriae probably to be a parasitoid of Drosophila malerkotliana Parshad & Paika. Some 
of the specimens from Zaire are paler and a few have pale apical antennal segments. 

SIRENES gen. n. 

Type species: Sirenes sinis sp. n. 

Diagnosis. 9 antenna 13-segmented, filiform-subclavate with from 3-8 segments with rhinaria, basal 
segments finely granulate; cf antenna 15-segmented, filiform-flagellate apically, segment 3 shorter or 
subequal to 4. Head viewed frontally longer than broad, eyes measured medially as far or further apart as 
height of an eye; weakly coriaceous in region of antennal sockets and eye margins; supra-clypeal area 
raised; face with sparse hairs; malar grooves weakly indicated with weak striations on either side; viewed 
dorsally ocellar area weakly coriaceous (Fig. 172), extending to occipital carina. Pronotal plate viewed 
dorsally rectangular (cf. Fig. 191), lateral fovea on either side weak and open; pronotum either side of 
pronotal plate densely pubescent. Mesoscutum polished medially, sometimes strongly coriaceous on 
lateral margins; notauli not indicated, parapsidal furrows or lateral lines present apically (Fig. 189). 
Mesopleuron polished, sometimes very weakly coriaceous, mesopleural suture or carina absent; sub- 
pleural area densely pubescent, antero-ventral cavity obscured by dense woolly pubescence enveloping the 
lateral margins of propodeum; metapleuron smooth and polished. Lateral bars of scutellum strongly 
striate; scutellar fovea large; scutellar disc coriaceous to weakly broken striate, almost square apically; 
scutellar cup (scutellar plate) long and narrow, rarely oval, not reaching apical margin of scutellar disc, 
apical fovea long and narrow. Propodeum completely covered dorsally and laterally with dense woolly 
pubescence. Segment 1 of gaster almost completely obscured by ring of dense woolly pubescence at base of 
tergite 2; tergites 2-5 (or less often 2-4) visible in lateral view, tergite 2 occupying most of visible area; apex 
of tergite 2 and visible parts of tergites 3-4 sculptured. Wing surfaces pubescent, margins ciliate, longest 
apically; fore wing with radial cell open or closed. Legs long, with scattered pubescence. 

Distribution. Afrotropical Region. 

Discussion. Sirenes is most closely related to Glauraspidia but the sculpture of the head, mesoscutum and 
mesopleuron together with the usually closed radial cell of the fore wing and the form of antenna in both 
male and female distinguish it. 

Key to the species of Sirenes gen. n. 

Females 

1 Antenna filiform, rhinaria present on the 8-9 apical segments, segment 3 shorter than 4 (4-13 

subequal in length) (Fig. 174); scutellar cup sharply declined apically (Fig. 175). spiosp. n.(p. 204) 

- Antenna weakly clavate , rhinaria present on a lesser number of apical segments, segments 3 and 

4 subequal in length; scutellar cup either weakly or sharply declined (Fig. 175,176) 2 

2 Strong rhinaria present on the 3 apical segments of antenna, segment 10 with weak rhinaria 

(Fig. 177); scutellar cup tear-shaped, sharply declined (Fig. 176); gaster finely punctate 
apically (Fig. 193); mesopleuron weakly coriaceous; radial cell of fore wing open on margin 
(Fig. 194) silenussp. n.(p. 203) 

- Strong rhinaria present on 4 or more apical segments of antenna; scutellar cup long and narrow 

or weakly oval apically; gaster either punctate or sculptured; mesopleuron sculptured or 
polished ; radial cell of fore wing closed 3 

3 Rhinaria present on the 4 apical segments of antenna, forming a weak but distinct club 

(Fig. 178); eye margins smooth and polished. 
Mesopleuron smooth and polished; gaster finely coriaceous apically (cf. Fig. 195). 

steropes sp. n. (p. 204) 



202 J. QUINLAN 

- Rhinaria present on 5 or more apical segments; outer lateral margins of eyes polished or 

sculptured 4 

4 Rhinaria present on 5 apical segments of antenna (Fig. 179) 5 

- Rhinaria present on more than 5 apical segments of antenna (Figs 17 1 , 1 80) 7 

5 Antenna moniliform, segment 3 as long as 4+5, 4-8 a little longer than broad (Fig. 179). 

Scutellar disc coriaceous to longitudinally striate (Fig. 189) orbilus sp. n. (p. 202) 

- Antenna weakly sub-clavate, segment 3 shorter than 4+5, 4-8 at least twice as long as broad 

(Fig. 181) 6 

6 Mesopleuron finely and densely coriaceous (Fig. 182); outer lateral margins of eyes coriaceous. 

syrtes sp. n. (p. 205) 

- Mesopleuron smooth and polished; outer lateral margins of eyes smooth and polished. 

floccussp. n. (p. 202) 

7 Rhinaria present on the 6 apical segments of antenna (Fig. 180); mesopleuron smooth and 

polished; gaster densely but finely coriaceous on apical half; scutellar cup declined 

(cf. Fig. 189) ; outer lateral margins of eyes smooth and polished syrinx sp. n. (p. 205) 

- Rhinaria present on the 7 apical segments of antenna; mesopleuron finely coriaceous 

(cf. Fig. 182); gaster coriaceous on apical half; scutellar cup weakly declined; outer lateral 
margins of eyes reticulate sinissp. n. (p. 203) 

Males 

At present males cannot be distinguished reliably. Although most species have males listed under material 
examined they have been associated mainly on the basis of collection data. The males tend to have 
differences in antennal lengths and shape as well as sculpture differences on the mesoscutum and 
mesopleuron. 

Sirenes floccus sp. n. 

(Figs 186-188) 

Description. $ antenna weakly subclavate, segment 3 shorter than 4+5, 4-13 subequal in length, 
segments 9-13 weakly swollen medially, with rhinaria, forming a weak club (Fig. 186). Head viewed 
frontally with eyes further apart, measured medially, than the height of an eye; frons and lateral areas of 
face smooth and polished; face and mandibular area with a few scattered hairs; occiput and vertex finely 
coriaceous; lateral margins of occipital carina with a line of dense pubescence. Pronotal plate with lateral 
fovea very small (cf. Fig. 191). Mesoscutum finely reticulate-coriaceous; lateral bars of scutellum 
coriaceous; scutellar cup almost reaching apex of scutellar disc, weakly rounded apically. Mesopleuron 
smooth and polished; anteroventral cavity hairless. Gaster with apex of tergite 2 reticulate; visible parts of 
3-5 finely sculptured (Fig. 187). Fore wing with radial cell closed, longer than wide, veins Rs + M and M 
indicated, weakly pigmented (Fig. 188). Colour: antenna dark brown; head, thorax and gaster chestnut 
brown; legs orange-yellow. 
CT unknown. 

Material examined 

Holotype $, Zaire: Massif Ruwenzori, Mont Ngulingopres. Nyamgaleke, 2500m, exI.N.A., 3.vi.l954 
(Vanschuytbroeck & Synave) (MR AC). 

Paratypes. Zaire: 5 $ (MRAC). 

Remarks. Separated from syrtes by mesopleural and eye margin differences. 

Sirenes orbilus sp. n. 

(Figs 179, 189, 190) 

Description. $ antenna moniliform, segment 3 subequal to 4+5, 4 longer than 5, 5-8 subequal, each 
slightly longer than wide, segments 9-13 forming a club, with rhinaria (Fig. 179); <$ antenna with segment 
3 shorter than 4, 4 longer than 5, 5-15 subequal, becoming thinner apically. Head viewed frontally with 
eyes further apart, measured medially, than the height of an eye ; frons and lateral areas of face smooth and 
polished; mandibular area with a few scattered hairs; occiput and vertex with fine reticulate-coriaceous 
sculpture; lateral margins of occipital carina with a line of dense pubescence. Pronotal plate with lateral 
fovea very small and open (this is difficult to appreciate) (cf. Fig. 191). Mesoscutum finely sculptured on 
lateral margins, polished medially (Fig. 189); lateral bars of scutellum striated, scutellar fovea smooth and 
polished; scutellar disc with coriaceous-reticulate-striate sculpture; scutellar cup narrow, tear drop- 



AFROTROPICAL EUCOILIDAE 203 

shaped, weakly declined apically (Fig. 189); apex of scutellar disc weakly rounded. Mesopleuron smooth 
and polished; lower margin of metapleuron densely pubescent, anteroventral cavity obscured by dense 
pubescence extending to lateral margin of propodeum. Segment 1 of gaster obscured from view by 
propodeal pubescence and base of tergite 2; apex of tergite 2 reticulate, visible parts of tergites 3-5 
reticulate (cf. Fig. 187); hypopygium inconspicuous. Wings narrow, rounded apically; radial cell of fore 
wing completely closed, almost triangular (Fig. 190); veins Rs + M and M not visible either as folds or 
pigmentation. Legs with femora and tibiae very weakly sculptured. Colour: antenna orange-yellow, apical 
segments dark; head, thorax and gaster chestnut brown; legs orange-yellow. 

Material examined 

Holotype $, Zaire: Rutshuru, 1285 m, ll.vii.1935 (de Witte) (MRAC). 
Paratypes. Zaire: 2 C? (MRAC). 

Remarks. Distinguished from other species in the genus by the moniliform antenna of the female (Fig. 179) 
and almost triangular radial cell of the fore wing. 

Sirenes silenus sp. n. 

(Figs 176, 177, 191-194) 

Description. 5 antenna weakly subclavate apically, segments 3-10 subequal in length, apical three 
segments with pronounced rhinaria, segment 10 with weak rhinaria, forming a weak club (Fig. 177). Head 
viewed frontally with eyes as far apart measured medially as the height of an eye ; face smooth and polished , 
reticulate on inner margins of eye, with a few scattered hairs; malar grooves percurrent; vertex and occiput 
reticulate-coriaceous; lateral margins of occipital carina with a line of dense pubescence. Pronotal plate 
rectangular, lateral fovea indistinct, appearing open (Fig. 191). Mesoscutum reticulate-coriaceous, 
impressions in place of notauli; lateral bars of scutellum coriaceous-striate; scutellar fovea deep, smooth 
and polished; scutellar cup long and narrow, dew drop-shaped, apical half oval and declined; scutellar disc 
squarish apically, surface dull, with trace of sculpture (Fig. 176). Mesopleuron finely reticulate-coriaceous; 
metapleural anteroventral cavity obscured by dense pubescence on lower margins of mesopleuron and 
lateral margins of propodeum. Segment 1 of gaster in form of narrow ring, obscured by presence on 
propodeum and the dense narrow ring of woolly pubescence at base of tergite 2 of gaster; tergite 2 
occupying whole of visible area of gaster in lateral view; apex of gaster punctate; hypopygium incon- 
spicuous (Fig. 193). Wing rounded apically; radial cell of fore wing partially open on wing margin; veins 
Rs + M and M indicated but not pigmented (Fig. 194). Legs with femora and tibiae finely coriaceous; 
trochanters of normal proportions. Colour: antenna orange-yellow basally, club segments darker; head 
and dorsal surface of mesoscutum blackish; mesopleuron and gaster orange-brown; legs orange-yellow. 
Cf unknown. 

Material examined 

Holotype $, Kenya: Nairobi, Karen, 1982 (Dewhurst) (BMNH). 
Paratype. Zaire: 1 $ (MRAC). 

Remarks. The antennal and scutellar cup characters separate silenus from all other species in the genus. 

Sirenes sinissp. n. 

(Figs 171-173, 195) 

Description. $ antenna subclavate, segment 3 longer than 4, 4-6 subequal in length, segments 7-13 with 
rhinaria, forming a weak 7-segmented club (Fig. 171); cf antenna flagelliform, segments 3-5 subequal in 
length, segment 3 weakly curved, 4 weakly swollen medially. Head viewed frontally with eyes as far apart 
measured medially as the height of an eye; frons and lateral areas finely reticulate-coriaceous; mandibular 
area with a few scattered hairs; occiput and vertex with dense fine reticulate-coriaceous sculpture; lateral 
margins of occipital carina pubescent. Pronotal plate viewed frontodorsally rectangular, surface 
coriaceous, lateral foveae very small and open (this is difficult to appreciate). Mesoscutum polished 
medially, laterally margins densely and finely reticulate-coriaceous (Fig. 173); lateral bars of scutellum 
striate; scutellar fovea smooth and polished; scutellar cup long and narrow, apical half weakly declined; 
scutellar disc weakly strigose, apex squarish. Mesopleuron finely coraceous; anteroventral cavity bare; 
lower margins of mesopleuron (mesosternum) densely pubescent. Segment 1 of gaster in form of a narrow 
ring, longer than wide, completely obscured by propodeal pubescence and a dense ring of pubescence at 
base of tergite 2; apical third of segment 2 and visible parts of 3-5 finely reticulate-coriaceous (Fig. 195); 
hypopygium broad apically (Fig. 195). Wings broad apically; radial cell of fore wing completely closed; 



204 J. QUINLAN 

Rs + M and M indicated, not pigmented. Legs with trochanters clearly longer than broad. Colour: antenna 
dark brown-black; head and thorax blackish brown; gaster chestnut-red; legs orange. 

Material examined 
Holotype $, Cameroun: Nkoemvon, viii.-ix.1980 {Jackson) (BMNH). 
Paratypes. Cameroun: 4 $, 1 cf (BMNH). Zaire: 8 $ (MRAC). 

Remarks. Apart from antennal differences the female is distinguished from other species on gastral 
characters. The males oispio and sinis cannot be separated and their identification has been by association 
with females. 

Sirenes spio sp. n. 

(Figs 174, 175,185,196) 

Description. £ antenna with the apical 8 or 9 segments with rhinaria, segment 3 shorter than 4, filiform, 
4-9 subequal in length, 10-13 with more strongly pronounced rhinaria (Fig. 174). c? antenna flagelliform. 
Head viewed frontally finely coriaceous; eyes further apart measured medially than the height of an eye; 
malar grooves percurrent, obscured by striae on either side; mandibular area with scattered hairs; vertex 
and occiput reticulate-coriaceous; lateral margins of occipital carina with a line of dense pubescence. 
Pronotal plate rectangular, broad based, lateral foveae open (Fig. 196). Mesoscutum finely and densely 
reticulate-coriaceous, lateral lines present apically; lateral bars of scutellum striate; scutellar foveae 
smooth and polished, deep; scutellar cup elliptical, apical third with a declined fovea; scutellar disc 
reticulate-coriaceous, dull, apex squarish (Fig. 175). Mesopleuron finely reticulate-coriaceous-striate; 
lower margins of mesopleuron (mesosternum) densely pubescent; anteroventral cavity obscured by dense 
pubescence extending from lateral margins of propodeum. Segment 1 of gaster in form of a crenulate ring 
as wide as long (longer than wide in male), obscured by thin ring of dense woolly pubescence at base of 
tergite 2; tergites 2-4 of gaster visible in lateral view; apical quarter of tergite 2 reticulate, segments 3 and 4 
punctate (cf. Fig. 187); hypopygium short, broad apically. Wings broad; radial cell of fore wing completely 
closed; veins Rs + M and M weakly pigmented. Legs with coxae, femora and tibiae coriaceous. Colour: 
antenna dark brownish yellow; head and thorax dark chestnut-brown; gaster orange-brown; legs orange- 
brown. 

Material examined 
Holotype $, Zaire: Kivu, Rutshuru, 1285 m, 12.vii.1935 (de Witte) (MRAC). 
Paratypes. Zaire: 1 $, 3 cT (MRAC). 

Rearks. The female is separated on gastral characters from sinis, although the males cannot be 
distinguished other than by association with females. 

Sirenes steropes sp. n. 

(Figs 178, 199) 

Description. § antenna weakly subclavate, segment 3 shorter than 4+5, 4 longer than 5, 5-9 subequal in 
length, 10-13 with rhinaria forming a weak 4-segmented club (Fig. 178) ; cf antenna flagelliform, segment 3 
weakly curved (cf. Fig. 183). Head viewed frontally with eyes further apart than the height of an eye 
measured medially; face smooth and polished; malar grooves obscured by striations on either side; 
mandibular area with few scattered hairs; occiput and vertex strongly coriaceous-strigose (Fig. 199). 
Pronotal plate viewed frontodorsally with lateral foveae open (cf . Fig. 191). Mesoscutum polished, notauli 
absent, finely coriaceous on lateral margins; lateral bars of scutellum striated; scutellar fovea large, smooth 
and polished; scutellar cup long and narrow with apical fovea weakly declined (cf. Fig. 189); scutellar disc 
striate, squarish apically. Mesopleuron smooth and polished; lower margins of mesopleuron densely 
pubescent; anteroventral cavity obscured by pubescence. Segment 1 of gaster completely obscured by 
pubescence of propodeum and hairy ring at base of tergite 2 of gaster; tergites 2-4 of gaster visible in lateral 
view; apex of tergite 2 and visible parts of 3 and 4 punctate (cf. Fig. 187); hypopygium not pronounced. 
Wings broad apically, with short widely spaced hairs on surface, apical margins with fringe of hairs long; 
radial cell of fore wing closed; veins Rs + M and M indicated, not pigmented (cf. Fig. 188). Legs with 
trochanters longer than broad, distinctive. Colour: antenna yellowish basally, apical segments darker; 
head and thorax dark brown; gaster orange-yellow; legs orange-brown. 

Material examined 

Holotype $ , Kenya: 15 mis NE. Kisumu nr Lake Victoria, xi.1979 (Croft) (BMNH). 

Paratypes. Kenya: 1 $ (BMNH). South Africa: 1 $ (BMNH). Zaire: 38 $ , 8 Cf (BMNH and MRAC). 

Remarks. Separated from other species by the antennal characters. 



AFROTROPICAL EUCOILIDAE 205 

Sirenes syrinx sp. n. 

(Figs 180, 183, 197) 

Description. $ antenna filiform, weakly subclavate, segment 3 shorter than 4+5, 4-7 subequal in length, 
8-13 with rhinaria forming a weak 6-segmented club (Fig. 180); cf antenna flagelliform, segment 3 weakly 
curved, longer than 4 (Fig. 183). Head viewed frontally with eyes slightly further apart than the height of an 
eye measured medially; inner orbits of face finely coriaceous; malar grooves obscured by striations on 
either side; mandibular area of face with scattered hairs; occiput strongly coriaceous, lateral margins of 
occipital area pubescent. Pronotal plate viewed frontodorsally with lateral fovea open (cf. Fig. 191). 
Mesoscutum polished, finely coriaceous on lateral margins, notauli absent; lateral bars of scutellum striate; 
scutellar fovea large, smooth and polished; scutellar cup long and narrow with apical fovea weakly 
declined; scutellar disc squarish apically, surface striated and partially coriaceous (cf. Fig. 175). Meso- 
pleuron finely and densely coriaceous; lower margins of mesopleuron densely pubescent, anteroventral 
cavity without pubescence. Segment 1 of gaster petiolate, narrow, a little longer than wide, partially 
obscured by a ring of dense pubescence at base of tergite 2; apical third and visible parts of 3-5 
reticulate-coriaceous; hypopygium not pronounced (cf. Fig. 187). Wings broad apically, rounded; veins 
Rs + M and M weakly indicated; radial cell of fore wing completely closed (Fig. 197). Legs with 
trochanters longer than broad, distinctive. Colour: antenna dark brownish yellow; head, thorax and gaster 
chestnut red; legs orange-yellow. 

Material examined 
Holotype $, Zaire: Rutshuru, riv. Kanzarue, 1200m, 16.vii.1935 {de Witte) (MRAC). 
Paratypes. Zaire: 23 2 > 6 Cf (MRAC). South Africa: 1 $ [this specimen is particularly pale] (BMNH). 

Remarks. In some specimens the colour is very much paler and the sculpture of the mesoscutum is very 
weak. Separated from other species by the antennal and mesopleural characters. 

Sirenes syrt essp. n. 

(Figs 181, 182, 184) 

Description. $ antenna subclavate, segment 3 weakly curved and subequal in length to 4, 4-8 equal in 
length, 9-13 forming a weak club (Fig. 181); cf antenna with segment 3 strongly curved, rhinaria 
pronounced (Fig. 184). Head viewed frontally with eyes further apart than the height of an eye measured 
medially; face reticulate-coriaceous; malar grooves percurrent; occiput and vertex with reticulate- 
coriaceous sculpture; lateral margins of occipital carina with a line of dense pubescence. Pronotal plate 
with lateral fovea appearing closed (cf. Fig. 191). Mesoscutum reticulate-coriaceous; lateral bars of 
scutellum striate; scutellar fovea smooth and polished, deep; scutellar cup long and narrow, apical half of 
cup and disc weakly declined; surface of scutellar disc weakly striated, apex squarish (cf. Fig. 189). 
Mesopleuron with fine coriaceous sculpture; anteroventral cavity obscured by dense pubescence. Segment 
1 of gaster in form of a ring as wide as long, partially obscured by ring of pubescence at base of tergite 2; 
apex of tergite 2 with reticulate-coriaceous sculpture extending to visible parts of 3-5; hypopygium not 
pronounced (cf. Fig. 193). Wings rounded apically, broad; radial cell of fore wing completely closed; veins 
Rs + M and M weakly pigmented (cf . Fig. 197) . Legs with hind femur and tibia finely reticulate-coriaceous ; 
trochanters not pronounced. Colour: antenna brownish basally, apical segments darker; head and thorax 
blackish; gaster chestnut red; legs dull orange-yellow. 

Material examined 

Holotype $, Zaire: Nyasheke (vole. Nyamuragira), 182 m, 14-26. vi. 1935 {de Witte) (MRAC). 
Paratypes. Zaire: 1 $, 1 cT (MRAC). 

Remarks. The mesopleural characters separate syrtes from floccus. 



Acknowledgements 

I thank my colleagues in the Hymenoptera Section for advice and criticism during the preparation of this 
paper. My thanks are extended to the Keeper of Entomology, Dr L. A. Mound, for allowing me to 
complete this paper after my retirement. 



206 J. QUINLAN 

References 

Ashmead, W. H. 1903. Classification of the gall-wasps and parasitic cynipoids, or the superfamily 

Cynipoidea II. Psyche 10: 59-73. 
Barbotin, F., Carton, Y. & Kelner-Pillault, S. 1979. Morphologie etbiologie de Cothonaspis (Cothonaspis) 

boulardi n. sp. parasite de drosophiles. Bulletin de la Societe Entomologique de France 84: 20-26. 
Benoit, P. L. G. 1956. Contributions a l'etude de la faune entomologique du Ruanda-Urandi (Mission P. 

Basilewsky 1953). GIX. Hymenoptera Cynipidae. Annates du Musee Royal du Congo Beige (Serie 8: 

Zool.) 51: 532-550. 
Bridwell, J. C. 1919. Descriptions of new species of hymenopterous parasites of muscoid Diptera with 

notes on their habits. Proceedings of the Hawaian Entomological Society 4: 166-179. 
Cameron, P. 1890. Monograph of British Phytophagous Hymenoptera 3: 274 pp. London. 
1904. Descriptions of new genera and species of Hymenoptera from Dunbrody, Cape Colony. 

Records of the Albany Museum 1 (3): 125-160. 

1910. On a new genus and species of parasitic Cynipidae (Eucoilinae) from Cape Colony. Entomolo- 



gist 43: 180. 
Dahlbom, A. G. 1842. Onychia och Callaspidia Tvenne for Skandinaviens. Fauna nya Insekt-Slagten, 

horande till Gallaple-Steklarnes naturliga grupp 16 pp. , 3 tables, 2 pis. Lund. 
Foerster, A. 1869. Ueber die Gallwespen. Verhandlungen der Zoologisch-Botanischen Gesellschaft in 

Wien 19: 327-370. 
Hartig, T. 1840. Uber die Damilie der Gallwespen, Zeitschrift fur Entomologie (Germar) 2: 176-209. 

1841. Naturgeschichte der Gallwespen. Zeitschrift fur Entomologie (Germar) 3: 322-358. 

Hedicke, H. 1913. Beitrage zur Kenntnis der Cynipiden (Hym.). V. Neue zoophage Cynipiden der 

indomalayischen Region. Deutsche Entomologische Zeitschrift 1913: 441-445. 
1930. Beitrage zur Kenntnis der Cynipiden (Hym.). XVII. Zur Synonymie der Anacharitinae, 

Eucoilinae, Figiten. Deutsche Entomologische Zeitschrift 1930: 74-76. 
Hellen, W. 1960. Die Eucoilen Finnlands. Fauna Fennica 9: 1-31. 
Kieffer, J. J. 1901. Revision des Eucoilines. Feuilles desJeunes Naturalistes 31: 158-176. 
1904. Les Cynipides: Eucoilinae. In Andre, E. , Species des Hymenopteres d' Europe et d'Algerie 7 (2): 

603-629. 
1910a. Serphidae, Cynipidae, Chalcidae, Evanidae und Stephanidae aus Aequatorial- Africa. Wis- 



senschaftliche Ergebnisse der Deutschen Zentral-Afrika Expedition 3 (2): 1-29. 

— 19106. Nouveaux Cynipides exotiques. Bolletino del Laboratorio Zoologica Portici 4: 329-342. 

— 1911. Hymenoptera Cynipidae. In The Percy Sladen Trust expedition etc -. Transactions of the 
Linnean Society of London (Ser. 2 Zool.) 14: 309-313. 

— 1913. Proctotrupidae, Cynipidae et Evaniidae. In Voyage de Ch. Allaud et R. Jeanel en Afrique 



Orientale (1911-1912) Resultats scientifiques. Hymenoptera 1: 1-198. 

1916. Neue Beitrage zur Kenntnis der Philippinischen Cynipen. Philippine Journal of Science 11D: 



279-287. 
Nordlander, G. 1976. Studies on Eucoilidae (Hym., Cynipoidea) I. A revision of the north-western 

European species of Cothonaspis Htg. with description of a new species and notes on some other genera. 

Entomologisk Tidskrift91: 65-77. 
1980. Revision of the genus Leptopilina Foerster, 1869, with notes on the status of some other genera 

(Hymenoptera. Cynipoidea: Eucoilidae). Entomologica Scandinavica 11: 428-452. 
1981. A review of the genus Trybiographa Foerster, 1869 (Hymenoptera, Cynipoidea: Eucoilidae). 



Entomologica Scandinavica 12: 381-402. 

1982. Systematics and phylogeny of an inter elated group of genera within the family Eucoilidae 



(Insecta: Hymenoptera, Cynipoidea) 32 pp. Stockholm. 
Quinlan, J. 1978. Hymenoptera, Cynipoidea, Eucoilidae. Handbooks for the identification of British 

insects 8 (7b): 1-58. 
1979. A revisionary classification of the Cynipoidea (Hymenoptera) of the Ethiopian Zoogeographi- 

cal Region. Aspicerinae (Figitidae) and Oberthuerellinae (Liopteridae). Bulletin of the British Museum 

(Natural History) (Entomology) 39: 85-133. 

1986. A key to the Afrotropical genera of Eucoilidae (Hymenoptera), with a revision of certain 



genera. Bulletin of the British Museum (Natural History) (Entomology) 52: 243-366. 
Risbec, J. 1956. Hymenopteres parasites du Cameroun (2 et 3 contributions). Bulletin de VInstitut Francais 

d' Afrique Noire 18: 97-164. 
Thomson, C. G. 1862. Forsok till uppsallning och beskrifning av Sveriges Figiter. Ofversigt af Kongl 

Vetenskaps-Akademiens Forhandlingar 18: 395-420. 



AFROTROPICAL EUCOILIDAE 



207 



Weld, L. 1930. Notes on types (Hymenoptera: Cynipidae). Proceedings of the Entomological Society of 
Washington 32: 137-144. 

1931. Additional notes on types with description of a new genus (Hymenoptera Cynipidae). 

Proceedings of the Entomological Society of Washington 33 (a) 220-227. 

1944. Descriptions of new Cynipidae including two new genera (Hymenoptera). Proceedings of the 

Entomological Society of Washington 46: 55-66. 
1952. Cynipoidea (Hymenoptera) J 905 -1950. 357 pp. Ann Arbor. 

1960. A new genus of Cynipoidea (Hymenoptera). Proceedings of the Entomological Society of 

Washington 62: 195-196. 

Westwood, J. O. 1833. Notice on the habits of a cynipoideous insect parasitic upon the rose louse (Aphis 
rosae); with descriptions of several other parasitic Hymenoptera. Magazine of Natural History 6: 
491-495. 



Index 



Principal references are in bold; synonyms are in italics. 



Aglaotoma 173, 189 
Amphiglyphosema 172, 173, 179 
Angustacorpa 171, 172, 173, 174 
apella 173, 192, 194 
Apis top hyza 173, 189 
apsus 173, 175 
ascia 173, 181, 182 
atraticeps 173, 192, 194 
Aulacidea 172 
Aylax 172 

bantia 173, 187 
Bothrochacis 172, 173, 176 
boulardi 173, 192, 195 

camerounensis 173, 188 
cantusl73, 181, 182 
capito 173, 181,183 
casca 173, 190 
clavipes 192, 193, 195 
Cothonaspis 172 

Daruna 171 

Diglyphosema 172, 173, 179 
Diranchis 173, 189 
Ditrupaspis 173, 176 

Emargc.171,173,174, 181 
erinna 173, 187, 188 
erythropoda 173, 177 
Eucoila 172, 173, 186 
Eucoilidae 173 
eupatorii 179 

fannius 173, 192, 193, 195 
faunus 173, 192, 193, 196 
Figitinae 172 



fimbriata 192, 193, 196 
floccus 173,202 

Ganaspis 171 
Glauraspidia 172, 173, 189 

heterotoma 173, 193, 196 

itys 173, 197 

Kleidotomal71,174 

latesulcatum 173, 179, 180 
laverna 173, 181, 183 
Leptopilina P2, 173, 191 
longipes 192, 193, 197 
Lytosema 173, 186 

mahensis 173, 195 
mahensis 173, 193, 198 
marina 173, 187, 188 
matius 173, 182, 183 
micipsa 173, 182, 184 
microptera 190 
misensus 173, 192, 193, 198 

Nordlanderia 172 
numa 173, 182, 184 

orbilusl73,202 

palloris 173, 182, 185 
peleus 173, 182, 185 
persa 173, 175 
pexus 173, 181, 185 
pisonis 173, 193, 194, 198 
prodicus 173, 175 



Psilodora 173, 186 
Psilodoropsis 173, 186 

Rhoptromeris 172 
rufiventris 173, 177, 178 

Satpictes 173, 176 
scyphus 173, 191 
semirufa 173 

serratepilosa 173, 177, 178 

silenusl73,201,202 

sinis 173,202,203 

Sirenes 171, 172, 173, 174,201 

spio 173,204 

Stentorceps 172 

stercoraria 173 

steropes 173,201,204 

Stirenocoela 173, 176 

striaticollis 1 73 

Sub ti lis 190 

syphax 173, 193, 199 

syrinx 173, 202, 205 

syrtes 173,202,205 

themis 173, 182, 186 
thetus 173, 192. 199 
Trichoplasta 172, 174 
triton 173, 176 
Trybliographa 171 

utica 173. 179, 180 

vacuna 173, 182, 186 
veleda 173. 187. 189 
vesta 173. 193.200 
victoriae 173, 193,200 

Zaeucoila 172 



208 



J. QUINLAN 




Figs 1-7 1 , 2, frontal view of head of (1) Angustacorpa sp. ; (2) Trichoplasta sp. 3 , basal segments of male 
antenna of Angustacorpa sp. 4, mesonotum of Angustacorpa apsus. 5, gaster of A. apsus. 6, 7, female 
antenna of (6) A. apsus; (7) A. triton. 



AFROTROPICAL EUCOILIDAE 



209 




Figs 8-16 8, 9, female antenna of (8) Angustacorpa prodicus; (9) A. persa. 10, mesonotum of A. persa. 
ll,pronotalplateof Aperstf. 12,gasterof A triton. 13, 14, radial cell [of (13) A. triton\(\A) A. apsus. 15, 
16, Bothrochacis erythropoda (15) female antenna; (16) scutellum. 



210 



J. QUINLAN 



^^W> 




Figs 17-21 17, 18, Bothrochacis erythropoda (17) pronotal plate; (18) female gaster. 19-21, female fore 
wing of (19) B. erythropoda; (20) Kleidotoma sp. ; (21) Emargo sp. 



AFROTROPICAL EUCOILIDAE 



211 




Figs 22-31 22, 23, female gaster of (22) Kleidotoma psiloides; (23) Emargo micipsa. 24, 25, radial cell of 
(24) Glauraspidia sp.; (25) Sirenes sp. 26-30, Bothrochacis serratepilosa (26) female antenna; (27) 
pronotal plate; (28) hind tibia; (29) radial cell; (30) scutellar cup. 31 , basal segments of female antenna of 
B. rufiventris. 



212 



J. QUINLAN 




Figs 32-40 32, 33, Bothrochacis serratepilosa (32) scutellum; (33) female gaster. 34, 35, B. rufiventris (34) 
lateral view of scutellum; (35) gaster. 36, 37, Diglyphosema latesulcatum (36) basal segment of male 
antenna; (37) pronotal plate. 38, radial cell of D. eupatorii. 39, 40, D. utica (39) gaster; (40) mesonotum. 



AFROTROPICAL EUCOILIDAE 



213 




Figs 41-49 41, 42, radial cell of (41) Diglyphosema utica; (42) D. latesulcatwn. 43, antenna of D. utica. 
44-46, D. eupatorii (44) pronotal plate; (45) basal segment of gaster; (46) mesonotum. 47-49. D. 
latesulcatwn (47) female antenna; (48) mesonotum; (49) basal segments of gaster. 



214 



J. QUINLAN 




Figs 50-57 50, female antenna of Emargo sp. 51, 52, Emargo ascia (51) mesonotum; (52) female gaster. 
53-57, female antenna of (53) E. laverna; (54) E. pexus; (55) E. capito; (56) E. cantus; (57) E. ascia. 



AFROTROPICAL EUCOILIDAE 



215 




Figs 58-66 58, female antenna of Emargo matius. 59, scutellum of E. palloris. 60, mesonotum of E. 
numa. 61-64, female antenna of (61) E. themis; (62) E. vacuna; (63) E. micipsa; (64) E. peleus. 65, radial 
cell of E. ascia. 66, mesonotum of E. cantus. 



216 



J. QUINLAN 




Figs 67-75 67, radial cell of Emargo cantus. 68, 69, mesonotum of (68) E. capito; (69) E. matius. 10,11., 
E. numa (70) female antenna; (71) mesonotum. 72, female antenna of E. palloris. 73, gaster of E. peleus. 
74, 75, mesonotum of (74) E. pexus; (75) E. eciton. 



AFROTROPICAL EUCOILIDAE 



217 




Figs 76-83 76-80, Eucoila crassinerva (76) female antenna; (77) head; (78) basal segments of male 
antenna; (79) mesonotum; (80) radial cell. 81, fore wing of Trybliographa rapae. 82, 83, female antenna 
of (82) Eucoila marina; (83) E. erinna. 



218 



J. QUINLAN 




Figs 84-91 84, 85, female antenna of (84) Eucoila bantia; (85) E. erinna. 86, 87, E. bantia (86) pronotal 
plate; (87) mesonotum. 88-90, E. marina (88) mesopleuron; (89) pronotal plate; (90) radial cell. 91, 
scutellum of E. erinna. 



AFROTROPICAL EUCOILIDAE 



219 




Figs 92-102 92, radial cell of Eucoila erinna. 93-96, Glauraspidia microptera (93) female antenna; (94) 
male antenna; (95) pronotal plate; (96) radial cell. 97, radial cell of G. scyphus. 98-102, G. casca (98) 
female antenna; (99) male antenna; (100) pronotal plate; (101) scutellum; (102) gaster. 



220 



J. QUINLAN 




Figs 103-11 103-105, Glauraspidia scyphus (103) female antenna; (104) basal segments of male antenna; 
(105) female gaster. 106-109, Leptopilina longipes (106) basal segments of male antenna; (107) 
mesonotum; (108) gaster; (109) hind coxae. 110, 111, mesonotum of (110) L. clavipes; (111) L. 
fimbriata. 



AFROTROPICAL EUCOILIDAE 



221 



.. ■ V ^, 



116 



u, 



113 



i 



114 




Figs 112-119 112, radial cell of Leptopilinathetus. 113-119, female antenna of (1 13) L.longipes; (114) L. 
misensus; (115) L. boulardi; (116) L. fimbriata; (117) L. thetus; (118) L. /fy5; (119) L. ape//a. 



222 



J. QUINLAN 




Figs 120-126 120, female antenna of Leptopilina fannius . 121, mesopleuron of L. boulardi. 122, radial 
cell of L. atraticeps. 123-125, scutellum of (123) L. atraticeps; (124) L. heterotoma; (125) L. victoriae. 
126, mesopleuron of L. heterotoma. 



<^ 



AFROTROPICAL EUCOILIDAE 



223 




130 



134 



Figs 127-135 127, 128, Leptopilina syphax {111) scutellum; (128) radial cell. 129, 130, L. pisonis (129) 
radial cell; (130) mesopleuron. 131, mesopleuron of L. victoriae. 132, female antenna of L. vesta. 133, 
male antenna of L. mahensls. 134, mesopleuron of L. syphax. 135, scutellum of L. fimbriata. 



224 



J. QUINLAN 




Figs 136-147 136, scutellum of Leptopilina clavipes . 137, 138, L.fimbriata (137) male antenna; (138) fore 
wing. 139, male antenna of L. longipes. 140, 141, radial cell of (140) L. heterotoma; (141) L. misensus. 
142, male antenna of L. syphax. 143, scutellum of L.fannius. 144, 145, male antenna of (144) L.fannius', 
(145) L. vesta. 146, 147, L. victoriae (146) radial cell; (147) male antenna. 



AFROTROPICAL EUCOILIDAE 



225 




Figs 148-159 148-150, Leptopilina apella (148) scutellum; (149) propodeum; (150) radial cell. 151, 152, 
pronotal plate of (151) L. atraticeps; (152) L. boulardi. 153, propodeum of L. boulardi. 154-156, L. 
fannius (154) metapleuron; (155) radial cell; (156) female antenna. 157, 158, L. faunus (157) meta- 
pleuron; (158) radial cell. 159, pronotal plate of L. apella. 



226 



J. QUINLAN 




Figs 160-170 160, female antenna of Leptopilina heterotoma. 161, 162, L. itys (161) propodeum; (162) 
scutellum. 163, 164, L. mahensis (163) propodeum; (164) radial cell. 165-167, L. misensus (165) male 
antenna; (166) pronotal plate; (167) metapleuron. 168, antenna of L. syphax. 169, scutellum of L. 
thetus. 170, radial cell of L. vesta. 



AFROTROPICAL EUCOILIDAE 

176 



227 




Figs 171-180 171-173, Sirenessinis (171) female antenna; (172) vertex; (173) scutellum. 174, 175, S. spio 
(174) female antenna; (175) scutellum. 176, 177, S. silenus (176) scutellum; (177) female antenna. 
178-180, female antenna of (178) S. steropes; (179) S. orbilus; (180) S. syrinx. 



228 



J. QUINLAN 




Figs 181-189 181, 182, Sirenes syrtes (181) female antenna; (182) mesopleuron. 183, 184, male antenna of 
(183) S. syrinx; (184) S. syrtes. 185, mesopleuron of 5. spio. 186-188, S.floccus (186) female antenna; 
(187) gaster; (188) radial cell. 189, mesonotum of S. orbilus. 



AFROTROPICAL EUCOILIDAE 



229 




Figs 190-199 190, radial cell of Sirenes orbilus. 191-194, S. silenus (191) pronotal plate; (192) mesono- 
tum; (193) gaster; (194) radial cell. 195, gaster of S. sinis. 196, pronotal plate of 5. spio. 197, 198, radial 
cell of (197) S. syrinx; (198) S. spio. 199, vertex of head of S. steropes. 



British Museum (Natural History) 

Milkweed butterflies: their cladistics and biology 

P. R. Ackery & R. I. Vane- Wright 

The Danainae, a subfamily of the Nymphalidae , contains only some 150 species, yet aspects of 
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genetics. The popularity of danaines among biologists can certainly be attributed to this 
combination, within one small group, of so many of the factors that make butterflies such an 
interesting group to study. The obvious need to place this wealth of biological data within an 
acceptable systematic framework provided the impetus for this volume. 

Started in 1976 within the conventions of evolution by natural selection and Hennig's 
phylogenetic systematics, the book is now largely about natural history (what the animals have 
and do, where they live and how they develop) and natural groups - as revealed by a form of 
analysis approaching that practised by the new school of 'transformed cladistics'. The authors 
have prepared a handbook that will appeal to a wide range of biologists, from museum 
taxonomists to field ecologists. 

425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. 
ISBN 565 00893 5. 1984. Price £50. 



Titles to be published in Volume 56 



The legume-feeding psyllids (Homoptera) of the west Palaearctic Region 

By I. D. Hodkinson & D. Hollis 

A review of the Malvales-feeding psy Hid family Carsidaridae (Homoptera) 

By D. Hollis 

The world genera of Rhadalinae (= Aplocneminae) (Coleoptera: Melyridae) 

By E. R. Peacock 

A revision of some Afrotropical genera of Eucoilidae (Hymenoptera) 

By J. Quinlan 



Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk 
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